Journal of Theoretical Biology 386 (2015) 147–158

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Journal of Theoretical Biology

journal homepage: www.elsevier.com/locate/yjtbi

Analytical formulae for computing dominance

from species-abundance distributions
Tak Fung a,n, Laura Villain b, Ryan A. Chisholm a,c
a
National University of Singapore, Department of Biological Sciences, 14 Science Drive 4, 117543 Singapore
b
Institut National des Sciences Appliquées de Lyon, 20 Avenue Albert Einstein, Villeurbanne cedex, 69621 Lyon, France
c
Smithsonian Tropical Research Institute, Balboa, Ancón, Republic of Panama

H I G H L I G H T S

Ecosystem stability and functioning critically dependent on species evenness.

Evenness quantified as proportion of species that dominate total abundance.

Formulae for dominance derived for different species-abundance distributions (SADs).

High dominance found for SADs with a log-series, lognormal or exponential form.

Formulae used to show how dominance increases with environmental variance.

art ic l e i nf o a b s t r a c t

Article history: The evenness of an ecological community affects ecosystem structure, functioning and stability, and has
Received 15 April 2015 implications for biodiversity conservation. In uneven communities, most species are rare while a few
Received in revised form dominant species drive ecosystem-level properties. In even communities, dominance is lower, with
6 August 2015
possibly many species playing key ecological roles. The dominance aspect of evenness can be measured
Accepted 12 September 2015
Available online 25 September 2015
as a decreasing function of the proportion of species required to make up a fixed fraction (e.g., half) of
individuals in a community. Here we sought general rules about dominance in ecological communities by
Keywords: linking dominance mathematically to the parameters of common theoretical species-abundance dis-
Biodiversity tributions (SADs). We found that if a community’s SAD was log-series or lognormal, then dominance was
Evenness
almost inevitably high, with fewer than 40% of species required to account for 90% of all individuals.
Gamma
Dominance for communities with an exponential SAD was lower but still typically high, with fewer than
Lognormal
Log-series 40% of species required to account for 70% of all individuals. In contrast, communities with a gamma SAD
only exhibited high dominance when the average species abundance was below a threshold of
approximately 100. Furthermore, we showed that exact values of dominance were highly scale-depen-
dent, exhibiting non-linear trends with changing average species abundance. We also applied our for-
mulae to SADs derived from a mechanistic community model to demonstrate how dominance can
increase with environmental variance. Overall, our study provides a rigorous basis for theoretical
explorations of the dynamics of dominance in ecological communities, and how this affects ecosystem
functioning and stability.
& 2015 Elsevier Ltd. All rights reserved.

1. Introduction sustained and ongoing loss of biodiversity (Chapin et al., 2000;

Human populations have caused massive and widespread
changes to the structure of natural communities (Vitousek et al.,
1997; Chapin et al., 2000; Steffen et al., 2007). This has resulted in
n
Corresponding author.
E-mail addresses: tfung2000@gmail.com (T. Fung),
villain.laura@gmail.com (L. Villain), ryan.chis@gmail.com (R.A. Chisholm).
Cardinale et al., 2012; Naeem et al., 2012). Since biodiversity is a
fundamental determinant of ecosystem functioning, its loss affects
functioning and associated ecosystem services (Cardinale et al.,
2012; Naeem et al., 2012). Research over the last few decades has
focused on the negative effects of species loss on functioning
(Balvanera et al., 2006; Cardinale et al., 2006, 2012; Naeem et al.,
2012), but the number of species is just one component of biodi-
versity. Another important component is species evenness, which
describes the relative abundances of species in a community.

http://dx.doi.org/10.1016/j.jtbi.2015.09.011
0022-5193/& 2015 Elsevier Ltd. All rights reserved.
148 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158
Evenness is arguably more directly related than richness to the
trait diversity on which the level of functioning depends (Reiss
et al., 2009). Recent experiments have demonstrated substantial
positive effects of evenness on biomass (Wilsey and Potvin, 2000;
Mattingly et al., 2007; Zhang et al., 2012; Orwin et al., 2014), water
retention (Orwin et al., 2014), resistance to invasion (Wilsey and
Polley, 2002) and nutrient retention after leaching (Orwin et al.,
2014). Moreover, evenness is expected to change quicker than
richness in response to human stressors, since changes in abun-
dance occur before species go extinct. Therefore, quantitative
assessments of the causes and consequences of changing evenness
are essential for a thorough understanding of ecosystem structure
and function.
One metric of evenness is dominance, which has been used to
quantify the degree to which the total abundance of a community
is dominated by a subset of species (e.g., Grime, 1998; Barker et al.,
2002; Binkley, 2004; Davidar et al., 2005; Hillebrand et al., 2008;
Dornelas et al., 2011; ter Steege et al., 2013). Dominant species
tend to have large effects on flows of energy and matter in an
ecological network (Hillebrand et al., 2008; Berlow et al., 2009). In
particular, empirical evidence suggests that in autotrophic and
suspension-feeding invertebrate assemblages, the contribution of
a species to functioning can be predicted largely by its biomass –
the “mass ratio hypothesis” (Grime, 1998; Davies et al., 2011).
Anthropogenic stressors such as targeted exploitation (e.g., stocks
of Atlantic cod, Gadus morhua; Lotze and Worm, 2009) may dis-
proportionately affect dominant species, resulting in heavy loss of
functioning (Gaston and Fuller, 2008). Ecosystems with high
dominance also typically have many rare species, posing chal-
lenges for biodiversity conservation.
Studies have quantified dominance for particular communities
(e.g., Grime, 1998; Barker et al., 2002; Binkley, 2004; Davidar et al.,
2005; Hillebrand et al., 2008; Dornelas et al., 2011; ter Steege et al.,
2013), but as yet there is no general theory for how dominance
emerges from standard ecological models. In this study, we first
derive mathematical formulae expressing dominance as a function
of the parameters of four standard species-abundance distributions
(SADs) – the log-series, exponential, gamma and lognormal. Spe-
cifically, for each SAD we derive formulae specifying dominance as a
decreasing function of the number of species accounting for a
proportion u of the total number of individuals (dominance is
defined in this way because fewer dominant species equates to
higher dominance). As u increases, more species are required for
dominance of numerical abundance in the community considered.
We examine four SAD forms that have been commonly studied and
fitted to empirical data: the log-series (Fisher et al., 1943; Hubbell,
2001), exponential (Cohen, 1968), gamma (Brian, 1953; Plotkin and
Muller-Landau, 2002; Forster and Warton, 2007) and lognormal
(Preston, 1948; Engen et al., 2002; Volkov et al., 2003; Forster and
Warton, 2007; Engen et al., 2011) forms. After deriving the dom-
inance formulae, we proceed to apply them to (i) gauge the levels of
dominance expected in natural communities under biologically
plausible scenarios; (ii) assess dominance in the metacommunity of
tree species found in Amazonia, using the data of ter Steege et al.
(2013); and (iii) quantify how dominance is expected to change
with increasing environmental variance, using the dynamic model
of Engen and Lande (1996a). These three applications of our for-
mulae highlight their three-fold utility: they allow dominance to be
quantified quickly for discrete SADs, they permit dominance to be
quantified for continuous SADs, and they can be mathematically
analyzed further to derive general biological insights that cannot be
obtained from simulations alone.
2. Methods
2.1. Deriving analytical formulae for calculating dominance: discrete
SADs
To derive a formula specifying dominance given an SAD for a
community, the minimum fraction of species (i.e., the most
abundant ones) required to account for a fraction u of the indivi-
duals in the community, denoted by pu , must first be computed.
We consider the case u Z 0:5, for which the minimum set of spe-
cies is called the set of dominant species, making up at least half of
all individuals. As pu decreases, fewer species are required to
account for a fraction u of all individuals, such that dominance
increases. Thus, dominance is defined as Du ¼ 1  pu .
To calculate pu , we introduce a quantity nu , which is defined as
the abundance of the least abundant dominant species. Three
simple examples are now given to illustrate clearly the meaning of
pu and nu , together with some difficulties with interpretation in
specific cases. First, consider a community with S ¼ 8 species and
abundances f1; 1; 3; 4; 4; 7; 8; 12g. In this example, p0:5 ¼ 0:25
because we can account for half of the individuals with just one
quarter of all species (i.e., the two most abundant ones). We also
have n0:5 ¼ 8, because species with abundance greater than or
equal to 8 comprise half of the community. Second, consider
another community with eight species but with abundances
f1; 4; 4; 4; 5; 5; 5; 8g. Here we have p0:5 ¼ 0:375 because we can
account for half of the individuals with the three most abundant
species. However, we note that the value of n0:5 is biologically not
well-defined here. It might seem logical to take n0:5 ¼ 5, but this is
problematic because to get 50% of all individuals we only want
two of three species with abundance 5. Thus, the quantity nu is
generally not as useful as pu , although it is necessary to calculate
nu in the derivation of pu . As a final simple example, consider yet
another community with eight species, this time with abundances
f1; 3; 4; 6; 6; 7; 8; 15g. In this case, the top three species are required
to reach at least 50% of individuals, but we actually overshoot and
get 60% of individuals. Such overshoots arise from the funda-
mental discreteness of individuals and pose problems of inter-
pretation. Thus, in our mathematical treatment, we will work in
the high-diversity, large-community size limit in which overshoots
are negligible, with the understanding that in practical applica-
tions, theoretical values for pu S, where S is the expected total
number of species, will need to be rounded to the nearest integer
to get a whole number of species.
We first detail a method for deriving exact formulae for pu and
hence Du for discrete SADs, in the high-diversity, large-community
size limit. Consider a discrete SAD defined by a probability dis-
tribution ψ ðnÞ, specifying the probability that a randomly selected
species will have abundance n. We have:
sðnÞ sðnÞ
ψ ðnÞ ¼ ¼ 1
P
; ð1Þ
S
sðnÞ
n¼1
where sðnÞ is the expected number of species with abundance n. To
obtain nu , the equation
P
1 P
1
nsðnÞ nψ ðnÞ
FðnÞ ¼ n ¼ m
¼n¼ m
¼u ð2Þ
P1 P
1
nsðnÞ nψ ðnÞ
n¼1 n¼1
can be solved analytically for m. Here, FðmÞ is the fraction of
individuals belonging to species with abundance greater than or
equal to m.
In general, the value of nu obtained would not be an integer, so
it is rounded up to the nearest integer, ⌈nu ⌉. Species with an
abundance equal to or greater than ⌈nu ⌉ are insufficient to make
 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158 149

up a fraction u of all individuals, so a proportion η of species with analytical tractability, which is why we used 1, 0 and 0 for the
abundance ⌈nu ⌉  1 are required. The quantity η satisfies three distributions, respectively. For the continuous exponential,
X
1 X
1 all three values of δ (0, 0.5 and 1) are examined since analytical
nψ ðnÞ þ ηð⌈nu ⌉  1Þψ ð⌈nu ⌉  1Þ ¼ u nψ ðnÞ: ð3Þ formulae for nu and pu can be derived for all three δ values.
n ¼ ⌈nu ⌉ n¼1

Rearranging gives 2.3. Applying the formulae to quantify the typical level of dominance
in ecological communities
P
1 P
1
u nψ ðnÞ  nψ ðnÞ
n¼1 n ¼ ⌈nu ⌉ The formulae derived using the methods described in Subsec-
η¼ : ð4Þ
ð⌈nu ⌉  1Þψ ð⌈nu ⌉  1Þ tion 2.1 and 2.2, for two types of discrete SAD and four types of
The quantity pu , which is the minimum fraction of species continuous SAD, are applied to quantify dominance across a broad
required to make up a fraction u of all individuals, can then be range of biologically possible scenarios. Specifically, parameter
calculated as values are specified for each type of SAD examined to give a broad
range of average species abundances ðJ=SÞ, from small to very
P
1 P
1
u nψ ðnÞ  nψ ðnÞ large, and then dominance values calculated for these parameter
X
1
n¼1 n ¼ ⌈nu ⌉ X
1
pu ¼ ηψ ð⌈nu ⌉  1Þ þ ψ ðnÞ ¼ þ ψ ðnÞ: values using the formulae. To facilitate comparison of dominance
⌈nu ⌉  1
n ¼ ⌈nu ⌉ n ¼ ⌈nu ⌉ values between different types of SAD, dominance for each type is
ð5Þ plotted against J=S, which is defined for all SADs. It is noted that
J=S for an SAD is equal to the mean of the underlying distribution;
The exact formula for dominance is then Du ¼ 1  pu . Usually,
this can be seen for discrete SADs because
the formula for pu cannot be simplified further because of the
ceiling function. However, simpler analytical approximations for P
1 P
1
nsðnÞ nψ ðnÞ X
1
pu can be derived by retaining the generally non-integer but J n¼1
¼ 1 ¼ n ¼11 ¼ nψ ðnÞ ð9Þ
analytical form of nu and then using S P P
sðnÞ ψ ðnÞ n¼1
X
1 n¼1 n¼1
pu  pu ¼ ψ ðnÞ: ð6Þ
and it can be seen for continuous SADS using analogous calcula-
n ¼ nu
tions. In addition, we mathematically analyze the dominance for-
We derive pu and pu according to Eqs. (5) and (6), respectively, mulae for the continuous log-series, continuous exponential and
when ψ ðnÞ follows a discrete log-series distribution and when it lognormal SADs to gain general insights into how they change
follows a discrete exponential distribution. We then derive Du and with J=S.
Du for these two distributions as 1 pu and 1  pu , respectively. For the discrete log-series SAD with parameter x, we examine
the parameter range ½4:65  10  10 ; 0:980, which gives J=S from
2.2. Deriving analytical formulae for calculating dominance: con- 1.01 (corresponding to the maximum x) to 108 (corresponding to
tinuous SADs the minimum x). 1.01 is close to the theoretical minimum of
1 whereas 108 is on the order of magnitude of the median number
Dominance formulae for continuous SADs can be derived ana- of individuals found for 207 dominant tree species in Amazonia,
logously to those for discrete SADs. For these distributions, nu can which covers an area of 6.29 million km2 (ter Steege et al., 2013).
take non-integer positive values and can be found by solving Thus, the range of J=S that we examine is a wide range encom-
R1 passing most values likely to be encountered in reality. The range
nψ ðnÞdn
GðmÞ ¼ Rn1¼ m ¼u ð7Þ of λ for the discrete exponential SAD is chosen in the same way as
n ¼ δ ψ ðnÞdn
n
½1:00  10  8 ; 4:62, with the maximum value corresponding to J=
for n, where ψ ðnÞ is the probability density corresponding to S ¼ 1:01 and the minimum value corresponding to J=S ¼ 108 . For
abundance n and δ is the lower limit of the domain of the corre- each of the two types of discrete SAD, we calculate exact and
sponding probability distribution. An analytical formula specifying approximate dominance values, Du and Dnu, respectively, over the
pu can then be derived using range of parameter values considered. The values of u (fraction of
Z 1 individuals accounted for by the dominant species) examined
pu ¼ ψ ðnÞdn; ð8Þ range from 0.5 to 0.9 in increments of 0.1. We also quantify the
n ¼ nu
error Du-Dnu for each combination of u and x or λ (i.e. each com-
allowing the corresponding dominance formula to be derived bination of u and J/S).
using Du ¼ 1 pu . We follow this method to derive dominance Parameter values for the continuous SADs are chosen using the
formulae for four continuous distributions that approximate dis- same rationale as for the discrete SADs. Thus, the range of x for the
crete SADs: the continuous log-series, exponential, gamma and continuous log-series SAD is chosen to be ½4:79  10  10 ; 1:00,
lognormal distributions. Values of Du for the continuous log-series corresponding to a range of J=S of ½1:01; 108 . Similarly, the ranges
and exponential distributions can be considered as approxima- of λ for the continuous exponential SAD are chosen to be
tions to the exact discrete cases because the distributions permit ½1:00  10  8 ; 0:990, ½1:00  10  8 ; 1:96 and ½1:00  10  8 ; 100 for
non-integer species abundances, which are not observed in reality. lower abundance limits of δ ¼ 0, 0.5 and 1 respectively, to corre-
Because the abundance n is assumed to be continuous, there are spond to the same range of J=S. For the gamma SAD with shape
conceptual issues with choosing δ, the lowest value of n for a and rate parameters α and β, it was found that Du is independent
continuous distribution. With δ ¼ 1, values between 0.5 and 1 are of β and hence scale (θ ¼ 1=β ). This is expected because the scale
omitted, but these values become 1 when rounded to the nearest parameter does not change the shape of the gamma distribution.
integer, so could be considered biologically meaningful. With However, J=S depends on both α and β. Therefore, we (i) calculate
δ ¼ 0, values between 0 and 0.5 are included, but these values are a range of β values from previous studies on plant and insect
rounded to 0, which is not biologically meaningful since it repre- communities (Brian, 1953; Plotkin and Muller-Landau, 2002; For-
sents species with no individuals. Thus, it may be better to use ster and Warton, 2007); (ii) calculate the minimum and maximum
δ ¼ 0:5 instead. However, using δ ¼ 0:5 for the continuous log- values of β and for each, calculate the corresponding range of α
series, gamma and lognormal distributions results in loss of that would give a range of J=S of ½1:01; 108 ; (iii) calculate values of
150 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158
Du corresponding to the two ranges of α found; and (iv) plot
curves of Du against J=S, one for each of the two ranges of α. The
minimum and maximum β values found were 0.000321 and
0.00761 respectively, with the corresponding ranges of α being
½0:000324; 32100 and ½0:00769; 761000 respectively. Similarly,
for the lognormal SAD with log-scale and shape parameters μ and
σ , it was found that Du is independent of μ, but that J=S depends
on both parameters. Therefore, we calculated the minimum and
maximum values of μ from previous studies on plant and bird
communities (Preston, 1948; Volkov et al., 2003; Forster and
Warton, 2007) as 1.39 and 3.47 respectively, and for each value
calculated a corresponding range of σ that would give a range of
J=S of ½32:2; 108 . Values of J=S lower than 32.2 are not considered
because they correspond to negative values of σ when the max-
imum value of μ ¼ 3:47 is used. For each range of σ , we calculate
corresponding values of Du and plot them against J=S. The values
of u examined range from 0.5 to 0.9 in increments of 0.1, as for the
discrete SADs. In addition, for each of the log-series and expo-
nential SADs, the error in Du arising from using the continuous
version is quantified as the difference between Du derived using
the exact formula for the discrete version minus Du derived using
the formula for the continuous version, for each combination of u
and J=S.
2.4. Applying the formulae to assess dominance for tree species in
Amazonia
To estimate the total number of tree species in Amazonia (the
Amazon and Guiana Shield), ter Steege et al. (2013) fitted a dis-
crete log-series SAD to species population abundances estimated
from their sample data and then extrapolated from this distribu-
tion. Using this method, they estimated that Amazonia has
approximately 16,000 tree species. However, their low sampling
intensity of 0.0002% (not 0.002% as ter Steege et al. state on p. 4 of
their Supplementary material) means that the distribution of rare
species is essentially unknown. Thus, other theoretical SADs, not
considered by ter Steege et al. (2013), provide equally good fits to
their estimated population abundances and give virtually the same
sample SAD (Supplementary material 1). These other theoretical
SADs lead to vastly different estimates of the total Amazon tree
species richness S. The examples in Supplementary material 1
show plausible Amazon-scale discrete log-series SADs truncated
such that S¼ 12,000, 9,000 and 6,000, with removal of increasingly
more of the least abundant species. To calculate corresponding
dominance values for these alternative distributions, we adapt our
dominance formulae for log-series SADs and apply them to the
truncated log-series SADs. This provides an idea of the range
within which the true dominance may vary. Dominance is
expected to decrease with greater truncation because the domi-
nant species would likely form a higher proportion of a smaller
total number of species.
Given the fundamental uncertainty in the distribution of rare
species in poorly sampled species-rich communities, a more
informative approach could be to fit only the observed portion of
the SAD, calculate the dominance of the fitted SAD and use it as an
estimate of a lower dominance bound. We apply this approach to
the Amazonian tree metacommunity by calculating dominance for
a log-series SAD fitted to the sample data and truncated to give
S  5; 000, corresponding approximately to the number of
observed species.
2.5. Applying the formulae to quantify how dominance changes with
environmental variance
In the dynamic community model of Engen and Lande (1996a),
the abundance ðn Z 1Þ of each species changes according to
dn dBðtÞ
¼ rn  ngðnÞ þ σ r ðnÞ ; ð10Þ
dt dt
where r is the per-capita growth rate, gðnÞ is the density-
dependent per-capita mortality rate, and BðtÞ is a Wiener process
representingstochastic  variation in the growth rate, with variance
σ 2r ðnÞ ¼ σ 2e þ σ 2d =n . This variance is the sum of two components,
the first representing environmental variance and the second
demographic variance. Under a Gompertz growth curve gðnÞ ¼ γ ln
ðn þ εÞ with ε ¼ σ 2d =σ 2e (there is a typo on p. 175 of Engen and Lande
(1996a) – they mistakenly stated that ε ¼ σ 2e =σ 2d ), the SAD at the
steady state is given by
"   2 #
A 1 lnðn þ εÞ  r=γ
SðnÞ ¼ pffiffiffiffiffiffiffiffiffiexp  ; ð11Þ
ðn þ εÞσ e π =γ 2 σ 2e =2γ
where A is a scaling constant (Engen and Lande, 1996a). Explicitly,
pffiffiffiffiffiffiffiffiffi (   )
2ω π =γ γ r 2
A¼ exp lnð1 þ ε Þ  ; ð12Þ
σe 2σe γ
where ω is the per-capita speciation rate. Using the transformation
z ¼ n þ ε, SðzÞ follows a truncated lognormal distribution
pffiffiffiffiffiwith
ffi log-
scale parameter μ ¼ r=γ and shape parameter σ ¼ σ e = 2γ over the
range ½1 þ ε; 1Þ. This distribution can be approximated by the full
lognormal distribution with range ½0; 1Þ if the density of the full
distribution is approximately zero in the range ½0; 1 þ εÞ, which is
the case for large communities. In this case, we apply our dom-
inance formula for the lognormal distribution to explicitly relate
dominance to environmental variance. We mathematically analyze
this relationship to show how dominance changes with increasing
environmental variance, and illustrate the trend found with
example parameter values.
3. Results
3.1. Analytical dominance formulae for SADs
For the six types of SAD that we considered, explicit formulae
were derived specifying dominance, measured as 1 minus the
minimum proportion of species required to account for a propor-
tion u of all individuals. This measure of dominance is denoted by
Du and varies between 0 and 1. In addition, for the two types of
discrete SADs, we derived formulae specifying Du , which is an
approximation to the exact dominance measure Du . Table 1 lists
the formulae obtained, together with formulae specifying the
probability distributions underlying the SADs. Appendix A pro-
vides details of how the formulae were derived following the
methods detailed in Subsection 2.1 and 2.2. It is seen that the
formulae for the (continuous) gamma and lognormal SADs depend
only on the shape parameter, that is, α for the gamma distribution
and σ for the lognormal distribution.
3.2. Quantifying dominance typically found in ecological
communities
In general, Du values obtained from the formula for the discrete
log-series were high, indicating that numerical abundance was
dominated by relatively few species (Fig. 1). When u increased
from 0.5 to 0.9 (in increments of 0.1), Du remained above 0.7 for
average species abundances (J=S) greater than 100. As J=S
 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158 151

Table 1
Formulae specifying the six types of species-abundance distributions (SADs) examined and the corresponding dominance. Each SAD is specified by a probability mass
function or probability density function (for discrete and continuous SADs, respectively), ψ ðnÞ, describing the probability or probability density of a species having abundance
n, respectively. Dominance is measured as Du ¼ 1 pu , where pu is the minimum proportion of species required to obtain a proportion u of all individuals. For each of the two
discrete SADs, an approximation to Du is also given as Du , which uses an approximation to pu that is specified by Eq. (6).

Probability distribution underlying SADa Dominance formula(e)

n h i
Discrete log-series ð1  xÞ
ψ ðnÞ ¼ nlog ; ð1  xÞ u  ð1  xÞ⌈nu ⌉  1 ⌈nu ⌉  1
ð1xÞ Du ¼ 1  log ð1=xÞ xð⌈nu ⌉  1Þ þ ð1  xÞ Φð1  x; 1; ⌈nu ⌉Þ ;
nZ1 log ðuÞ
nu ¼ 1 þ
log ð1  xÞ

uð1  xÞ log ðuÞ

Du ¼ 1  Φ 1  x; 1; 1 þ
log ð1=xÞ log ð1  xÞ

  h i
Discrete exponential ψ ðnÞ ¼ eλ  1 e  λn ; u  e  λ⌈nu ⌉ ð1  ⌈nu ⌉ þ ⌈nu ⌉eλ Þ
Du ¼ 1  eλ ðeλ  1Þð⌈nu ⌉  1Þ þ e  λ⌈nu ⌉ ;
nZ1 1 W  1 ðuye  y Þ
nu ¼  ;
1  eλ λ
λ
y¼
1 eλ   
Du ¼ 1  exp λ  yþ W  1 uye  y

n
Continuous log-series ψ ðnÞ ¼ nΓð0; ð1  xÞ
 log ð1  xÞÞ
Du ¼ 1  liðliuð1  xÞÞ
ð1  xÞ
,
nZ1

h  i
Continuous exponential ψ ðnÞ ¼ λeλð  n þ δÞ ; Du ¼ 1  exp 1 þ δλ þ W  1  uð1 þ δλÞ
eð1 þ δλÞ
nZδ

Gamma (continuous) α
nα  1 e  βn Γðα; Q  1 ð1 þ α;uÞÞ
ψ ðnÞ ¼ β ΓðαÞ ; Du ¼ 1  ΓðαÞ
nZ0


Lognormal (continuous)
ψ ðnÞ ¼
a
Special functions used in the SAD and dominance formulae are the Gamma function ΓðzÞ, the incomplete gamma function Γða; zÞ, the Lerch transcendent Φðz; s; aÞ, the
Lambert W function W  1 ðzÞ, the inverse regularized gamma function Q  1 ða; zÞ, the complementary error function erfcðzÞ, the inverse complementary error function
1
erfc ðzÞ, and the logarithmic integral liðzÞ (Abramowitz and Stegun, 1972).
decreased below 100, Du decreased non-linearly to low values,
dropping below 0.2 when u ¼ 0:9 (Fig. 1). Values of Du for the
discrete exponential were smaller than for the discrete log-series
(Fig. 1), indicating that numerical abundance was dominated by
more species. Nonetheless, Du was generally quite high: when u
increased from 0.5 to 0.7, it remained above 0.6 for J=S 4 100
(Fig. 1a–c), only dropping below 0.6 for J=S 4 100 when u
increased further to 0.8 and 0.9 (Fig. 1d and e). As for the discrete
log-series, Du for the discrete exponential decreased non-linearly
when J=S decreased below 100, to below 0.2 when u ¼ 0:9 (Fig. 1).
The Du values for both the discrete log-series and exponential
were found to be very close approximations of the exact dom-
inance values (Du values) (Fig. 2). The absolute error between the
approximate and exact values was almost zero except for very low
values of J=S below 10, when the absolute error increased but still
remained below 0.1 (Fig. 3). Therefore, trends in Du closely fol-
lowed trends in Du .
Values of Du for the continuous log-series are close approx-
imations of the corresponding values for the discrete log-series
(Fig. 2a), with absolute errors typically o 0.05 and always o 0.11
(Fig. 3a). Similarly, values of Du for the continuous exponential
were typically close approximations of corresponding values for
the discrete exponential, regardless of whether a lower limit of
δ ¼ 0, 0.5 or 1 was used (Figs. 2b and S2 in Supplementary
material 2); absolute errors were virtually zero for J=S above 100
(Figs. 3b and S3 in Supplementary material 2). The absolute
errors were typically higher as J=S decreased below 100, reaching
values close to 0.4 with δ ¼ 0 (Figs. 3b and S2). However, with
δ ¼ 0:5, the errors remained virtually zero down to J=S ¼ 10 and
remained below 0.2 for lower J=S values (Fig. 3b), and with δ ¼ 1,
the errors remained below 0.1 as J=S decreased below 100 (Fig.
exp
2
 ðμ  log2ðnÞÞ
1
Du ¼ 1  12 erfc pσffiffi2  erfc ð2ð1  uÞÞ
pffiffiffiffi ;
2σ
nσ 2π
nZ0
S3). Thus, dominance trends for continuous log-series and
exponential SADs closely follow those for their discrete coun-
terparts. In addition, we showed
  analytically that for the con-
tinuous log-series, dDu =d J=S 40 for small and large J=S
(Appendix B). These analytical results hold for all u, thus gen-
eralizing the trends found from simulations for particular values
of u (Fig. 1). Furthermore, we proved that for the continuous

exponential with lower abundance limit δ 4 0, dDu =d J=S 4 0 for
all u (Appendix B), again generalizing the simulated dominance
trends (Fig. 1).
Values of Du for the lognormal distribution followed a similar
trend to those for the log-series and exponential, exhibiting high
values greater than 0.6 for J=S above 200, regardless of the value of
u tested, and decreasing non-linearly as J=S decreased below 200
(Fig. 1). For large J=S above 10,000, Du reached values close to 1 for
all values of u tested, exceeding Du for both the log-series and
exponential (Fig. 1). Values of Du for the gamma distribution
showed the opposite trend to that of the other three distributions,
decreasing rather than increasing with J=S (Fig. 1). For small J=S
below 50, Du attained high values above 0.6, but as J=S increased
above 50, Du decreased according to a non-linear sigmoidal shape,
to values below 0.6 (Fig. 1). We mathematical analyzed the form of
 
Du for the lognormal and found that dDu =d J=S 4 0 for all u
(Appendix B).
3.3. Assessing dominance of tree species in Amazonia
In their study of tree species in Amazonia, ter Steege et al.
(2013) fitted a discrete log-series SAD with a parameter value of
x ¼ 1:93  10  9 to species abundances estimated from sampled
data. This parameter value corresponds to J=S ¼ 2:58  107 and
152 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158

Fig. 1. Dominance values ðDu Þ for the discrete log-series, discrete exponential, (continuous) gamma and (continuous) lognormal SADs, calculated using the formulae in
Table 1. Panels (a)–(e) show how dominance changes across average species abundance (total number of individuals of all species, J, divided by total number of species, S) for
five different values of u, which is the fraction of individuals accounted for by the set of dominant species.

S  16; 000. Application of our exact dominance formula (Table 1) and

shows that for this SAD, 1.89% of species is required to account for  
uΦ 1  x; 1; nmin þ loglogð1ðuÞ
50% of all tree individuals, i.e. Du ¼ 0:981 for u ¼ 0:5. However, a Du ¼ 1
 xÞ
ð14Þ
truncated discrete log-series fits the data just as well (Supple- Φð1  x; 1; nmin Þ
mentary material 1). Following the method used to derive the
exact and approximate dominance formulae for the full log-series, respectively. In these formulae, nmin is the species abundance at
which the SAD is truncated, such that species with a lower abun-
we derived corresponding formulae for a truncated log-series as  
dance are removed; also, in (13), nu ¼ nmin þ log ðuÞ=log ð1  xÞ . The
" #
1 u  ð1  xÞ⌈nu ⌉  nmin dominance formula for the continuous version of the truncated log-
Du ¼ 1  þ ð1  xÞ⌈nu ⌉  nmin Φð1  x; 1; ⌈nu ⌉Þ
Φð1  x; 1; nmin Þ xð⌈nu ⌉  1Þ series can be derived analogously to that for the continuous log-
ð13Þ
 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158 153

Fig. 2. (a) Dominance values for the discrete log-series SAD using the exact and approximate formulae, compared with dominance values for the continuous log-series SAD
(formulae shown in Table 1). The panel shows how dominance changes across average species abundance (total number of individuals of all species, J, divided by total
number of species, S) for different values of u, which is the fraction of individuals accounted for by the set of dominant species. (b) is the same as (a) but for the discrete
exponential SAD and the continuous exponential SAD with a lower limit of δ ¼ 0:5.

Fig. 3. Errors in approximating the exact dominance values for (a) the discrete log-series SAD and (b) the discrete exponential SAD when using the approximate formulae
and the formulae for the continuous versions (formulae shown in Table 1). An error is calculated as the exact value minus the approximate value. Each panel shows how the
errors change across average species abundance (total number of individuals of all species, J, divided by total number of species, S) for different values of u, which is the
fraction of individuals accounted for by the set of dominant species. In (b), the continuous exponential SAD has a lower limit of δ ¼ 0.5.

series, as (13) and (14) because evaluation of the Lerch transcendent is slow
  with small x.
li uð1  xÞnmin
Du ¼ 1    : ð15Þ A lower bound for the dominance of tree species in Amazonia
li ð1  xÞnmin
was estimated by applying dominance formulae (13)–(15) with
Application of (13) with u ¼ 0:5 and nmin ¼ 84, 3,646 and nmin ¼ 550; 755, corresponding to S  5; 000, where 5000 is the
157,027, corresponding to S  12; 000, 9,000 and 6,000 respec- approximate number of observed species. All three formulae gave
tively, gave Du ¼ 0:975, 0.966 and 0.949 respectively. Application Du ¼ 0:940 to 3 decimal places. By plotting Du against nmin from
of (14) and (15) gave dominance values that are the same to 1 to 550,755, we confirmed our expectation that Du decreases with
3 decimal places; however, application of (15) is much faster than nmin , such that Du ¼ 0:940 can be taken as a lower bound.
154 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158

Fig. 4. (a) Lognormal distributions underlying (stationary) SADs produced by the dynamic community model by Engen and Lande (1996a), for three different values of the
square root of environmental variance (environmental standard deviation), σ e (measured in units of yr  1/2, not shown on graphs to improve their clarity). Values of the other
parameters are r ¼ 0:16yr  1 , γ ¼ 0:02 yr  1 , σ d ¼ 0:447 yr  1=2 and ω ¼ 10  4 yr  1 (for parameter definitions, see Subsection 2.5). (b) Dominance values for the lognormal
SADs, as calculated by applying the formula in Table 1 for five different values of u (fraction of individuals accounted for ffiffiffiffiffi the set of dominant species) and σ e ranging from
pby
0.224–0.380 yr  1/2. The log-scale parameter for the SADs is μ ¼ r=γ ¼ 8 and the shape parameter varied from σ ¼ σ e = 2γ ¼ 1:12 to 1.90.

3.4. Quantifying how dominance changes with increasing environ-
mental variance
We applied the dominance formula for the lognormal SAD
(Table 1) to the lognormal SADs arising from the dynamic com-
munity model by Engen and Lande (1996a) (as described in
Subsection 2.5). According to this formula, dominance increases
with the shape parameter σ , and hence the average species
abundance J=S (Subsection 3.2). SADs from the dynamic model
have a shape parameter that is proportional to σ e , which is the
square root of environmental variance. Thus, application of the
formula shows that dominance increases with σ e in the model.
The relationship between dominance and environmental variance
is illustrated in Fig. 4 for example parameterpsets ffiffiffiffiffiffiffiffiffithat
ffi are the same
except for σ e increasing by a factor of 1.7 from 0:05 ¼ 0:224yr  1=2 to
0.380 yr  1/2. This gave a range of σ 2e values that are in accordance
with empirical values collated by Lande et al. (2003; Table 1.2), typi-
cally on the order of 0.01–0.1 yr  1. Similarly, demographic variance
σ 2d was set at 0.2, giving a ratio ε ¼ σ 2d =σ 2e from 1.38–4, in accordance
with typical empirical ratios of 1 to 10 (Table 1.2 of Lande et al., 2003).
The dynamic model explicitly models speciation, which operates over
large spatial scales. Thus, values of the other parameters were chosen
to give large values of J and J=S, commensurate with communities at
large spatial scales. These quantities together with S can be calculated
using Eqs. (11) and (12), after translation of the abundance variable
and assuming that the transformed variable starts at zero (a reason-
able assumption for large communities). For the example shown in
Fig. 4, dominance increased with σ e by 12–68% for values of u ranging
from 0.5 to 0.9 in increments of 0.1. The sharpest increase was seen for
the highest u tested. In addition, as σ e increased, S decreased from
139,000 to 7.52; J decreased from 7:76  108 to 136,000; and J=S
increased from 5570 to 18,100.
4. Discussion
In this study, we examined four standard forms of SADs that
together have commonly been fitted to empirical data (e.g., Fisher
et al., 1943; Preston, 1948; Brian, 1953; Cohen, 1968; Hubbell,
2001; Engen et al., 2002; Plotkin and Muller-Landau, 2002; Volkov
et al., 2003; Forster and Warton, 2007; Engen et al., 2011) and
successfully derived analytical formulae quantifying dominance
according to Du ¼ 1  pu , where pu is the minimum proportion of
species required to account for a proportion u of all individuals
(Table 1). We also examined the discrete broken-stick (MacArthur,
1957) and Zipf-Mandelbrot (Frontier, 1985) SADs, but due to the
complexity of the SAD formulae, we were unable to derive cor-
responding analytical dominance formulae. However, in the limit
of large S, the broken-stick SAD converges to a gamma distribution
(Longuet-Higgins, 1971), which we examined (Table 1). The dom-
inance formulae derived facilitate quantitative investigation of
dominance by speeding up calculation of dominance for discrete
SADs (compared with simple summation, the formulae can be
orders of magnitude quicker); allowing dominance to be calcu-
lated for continuous SADs; and permitting further mathematical
analyses that yield general insights into how dominance changes
with key ecological variables. We also derived analytical formulae
to approximate Du for the discrete log-series and exponential
SADs, and found that these approximations gave almost zero error
when the average number of individuals per species was greater
than about 10. In addition, under these circumstances, dominance
formulae for the continuous versions of the SADs gave values that
were very similar to those from the exact formulae for the discrete
versions. Thus, the formulae for the continuous versions can be
used to give accurate estimates of dominance for any natural
community of reasonable size that is distributed according to one
of these two discrete SADs. The errors when using the approx-
imate formulae for the discrete SADs were typically lower than
those when formulae for the continuous SADs were used (Fig. 3).
This result is intuitive because the continuous SADs assume that
species abundance changes continuously, whereas in reality it is
discrete. However, despite this continuity assumption, the result-
ing continuous SADs largely retain the dominance structure of
their discrete counterparts (Fig. 2). Given lower errors, the discrete
formulae would be preferred in practice, except for the discrete
log-series SAD with low values of the parameter x. In this case, the
approximate discrete formula is slow to compute because of the
Lerch transcendent function, and so the continuous formula would
be preferred.
A striking result from application of our formulae is that
dominance was typically very high for reasonably large commu-
nities with individuals distributed according to a log-series,
exponential or lognormal distribution. When the average abun-
dance of a species exceeded 200 individuals, fewer than 40% of
species were required to account for 90% of all individuals for the
log-series or lognormal case (Du 40:6 for J=S 4 200 and u ¼ 0:9),
and 70% of all individuals for the exponential case (Du 4 0:6 for
J=S 4 200 and u ¼ 0:7). Given that many empirical SADs can be
 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158
fitted by these three distributions (Fisher et al., 1943; Preston,
1948; Cohen 1968; Hubbell, 2001; Engen et al., 2002; Volkov et al.,
2003; Forster and Warton, 2007; Engen et al., 2011; the lognormal
distribution is typically mixed with a Poisson sampling distribu-
tion), the concentration of abundance in so few species that we
typically found has important implications for community func-
tioning and stability. Communities with high dominance attain a
higher level of functioning if there is a positive correlation
between numerical dominance of a species and its level of func-
tioning (Hillebrand et al., 2008) – this is the classic positive
selection effect that has been demonstrated to be an important
mechanism underlying positive relationships between richness
and functioning for plant and invertebrate communities (Huston,
1997; Wilsey and Potvin, 2000; Hooper et al., 2005; Larsen et al.,
2005; Orwin et al., 2014). However, another implication of high
dominance is a decrease in the strength of multispecies processes,
thus reducing the prevalence of niche complementarity and
facilitation, both of which are important drivers of functioning
(Loreau and Hector, 2001; Cardinale et al., 2002; Hooper et al.,
2005; Carey and Wahl, 2010). In contrast to the selection effect,
this acts to decrease community functioning. Therefore, the net
effect of higher dominance depends on the balance of two coun-
tervailing forces – the selection effect and weaker multispecies
processes. A meta-analysis of 54 studies suggests that the selection
effect is typically stronger in forest communities, resulting in
greater community functioning with higher dominance (Zhang
et al., 2012). In addition to these effects on functioning, an increase
in dominance results in an increase in intraspecific and corre-
sponding decrease in interspecific interaction strengths, causing
changes in community stability. Studies on model and empirical
food webs suggest that weak interspecific interactions can
increase stability (May, 1972; McCann et al., 1998; Neutel et al.,
2002; Emmerson and Yearsley, 2004), such that higher dominance
is expected to result in more stable communities.
In contrast to the log-series, exponential and lognormal dis-
tributions, dominance values for the gamma distribution were
typically high only for communities with small average species
abundances. When the average species abundance was below 50
individuals, fewer than 40% of species were required to account for
90% of all individuals (Du 40:6 for J=S o 50 and u ¼ 0:9). However,
as the average species abundance increases beyond 50 individuals,
dominance decreases non-linearly, typically reaching values below
0.5. Thus, our formulae give opposite trends for how dominance in
gamma- and lognormally-distributed communities scale with
community size. This result is surprising given that these two
distributions can have similar shapes. However, it can be explained
by the tail of the lognormal distribution becoming heavier as the
value of the shape parameter, which correlates with community
size, increases, whereas the tail of the gamma distribution
becomes less heavy. Therefore, for communities that are fit well by
a gamma distribution (typically mixed with a Poisson sampling
distribution; e.g., Brian, 1953), it is advisable to rigorously test
whether a lognormal distribution can give a better fit or not, since
this can have important implications for determining dominance
and assessing its community effects. On the other hand, this is
difficult given that statistical estimates of the rare species tail of an
SAD are highly uncertain unless a community is intensively sam-
pled. This is also evident from our analysis of the SAD underlying
the tree metacommunity in Amazonia (discussed below), and
together these results emphasize the sensitivity of dominance to
sampling uncertainty in general.
A recent assessment of dominance among trees in Amazonia
involved fitting a discrete log-series SAD to sample plot data and
concluded that dominance is very high, in the sense that very few
species are required to account for 50% of all individuals (ter
Steege et al., 2013). We now see, from application of our analytical
155
formula for discrete log-series SADs, that this result follows
inevitably from fitting a log-series: the formula predicts high
dominance for values of the parameter x corresponding to large
communities with high average species abundances ðJ=SÞ. How-
ever, whether the actual distribution of Amazon trees is a log-
series is highly uncertain, given a low sampling intensity of
0.0002% (ter Steege et al., 2013). Alternative types of SADs can be
fitted to the Amazonian data equally well, and one of these is the
truncated discrete log-series (Supplementary material 1). We
adapted our calculations to derive dominance formulae for a
truncated log-series and used it to show that dominance in the
Amazonian tree flora can vary between 0.940 and 0.981, i.e., the
percentage of species making up 50% of individuals could be
anywhere between 1.9% and 6.0%. Thus, dominance is high, but the
exact degree of dominance currently remains uncertain. The true
dominance may even exceed the upper limit of 0.981 if there are
more rare species than predicted by a log-series. Furthermore, the
true level of dominance is probably fundamentally unknowable
from small sample plots, because it is not possible to calculate an
unbiased estimate of the total number of species from a small
sample (Gotelli and Chao, 2013; Supplementary material 1). A
more justifiable approach in such situations may thus be to cal-
culate dominance from a fitted distribution that covers only the
empirical range of species abundances and use it as an estimate of
a lower bound for dominance – for the Amazonian tree flora, such
an estimate would be 0.940.
We also applied our dominance formula for lognormal SADs to
the dynamic community model by Engen and Lande (1996a),
which predicts lognormal SADs as a function of parameters
representing the ecological mechanisms modeled. This allowed a
quantitative, process-based assessment of how dominance is
predicted to change with variations in the ecological mechanisms.
In our application, we focused on how dominance is expected to
change with environmental variance, which is modeled as sto-
chastic fluctuations in the growth rate. We showed analytically
that the model predicts increases in dominance with the degree of
environmental variance, and illustrated this with an example
whereby dominance increased by up to 68%. It is also possible to
show analytically that the model predicts a decline in species
richness with environmental variance, which was the case in the
example shown. Therefore, the model predicts that greater
environmental fluctuations homogenize communities by causing
extinctions and promoting dominance of abundance by pro-
portionally fewer species, in effect pushing species abundances
towards the extremes (Fig. 4a). This prediction is particularly
relevant in the context of forecasted increases in environmental
variance driven by climatic change, for example greater intra-
annual variation in precipitation resulting in more severe floods
and droughts (Knapp et al., 2008). Our predictions complement
previous theoretical predictions on how environmental variance
can increase extinction risk and decrease species richness (e.g.,
Adler and Drake, 2008; Ovaskainen and Meerson, 2010). Impor-
tantly, our predictions can be tested quantitatively using empirical
data in future studies. In addition, the homogenizing effect of
environmental variance has possible knock-on effects for func-
tioning and stability, as discussed above. This can be further
investigated in future studies by explicitly quantifying functioning
and stability using model outputs and possibly other theory. Fur-
thermore, future studies could use our dominance formulae in
conjunction with other stochastic community models (e.g., Dennis
and Costantino, 1988; Engen and Lande, 1996b; Hubbell, 2001) to
further investigate the mechanistic links between evenness and
ecosystem functioning and stability. Predictions from these studies
would facilitate understanding of the ongoing effects of anthro-
pogenically driven biodiversity loss (Cardinale et al., 2012), vital
for conservation and sustainable management.
156 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158

 
Acknowledgments X
1
eλ  λ⌈nu ⌉ 1  ⌈nu ⌉ þ ⌈nu ⌉eλ
nψ ðnÞ ¼ ; ðA:11Þ
n ¼ ⌈nu ⌉
eλ 1
We would like to thank two anonymous reviewers, whose
comments have helped to substantially improve the clarity and and
content of our work. TF and RAC are supported by the National X
1
University of Singapore, Singapore start-up Grant WBS R-154-000- ψ ðnÞ ¼ eλ  λ⌈nu ⌉ : ðA:12Þ
551-133. n ¼ ⌈nu ⌉

(A.10) and (A.11) can be derived using the identity

P1 2
k ¼ a kz ¼ ½z ða þ ð1  aÞzÞ=ð1  zÞ , which holds for 0 o z o 1.
k a
Appendix A. Details on derivation of dominance formulae
Thus,
 
This Appendix contains details of how the dominance formulae ueλ  eλ  λ⌈nu ⌉ 1  ⌈nu ⌉ þ ⌈nu ⌉eλ
pu ¼   þ eλ  λ⌈nu ⌉
in Table 1 were derived. eλ  1 ð⌈nu ⌉  1Þ
"   #
Discrete log-series λ u  e  λ⌈nu ⌉ 1  ⌈nu ⌉ þ ⌈nu ⌉eλ  λ ⌈n ⌉
¼e   þe u
: ðA:13Þ
eλ  1 ð⌈nu ⌉ 1Þ
In this case, FðmÞ ¼ u is
Using Du ¼ 1  pu gives the formula for Du in Table 1. Also,
ð1  xÞm  1 ¼ u; ðA:1Þ
X
1   
which can be solved for m to give pu ¼ ψ ðnÞ ¼ eλð1  nu Þ ¼ exp λ  y þ W  1 uye  y : ðA:14Þ
n ¼ nu
log ðuÞ
nu ¼ 1 þ : ðA:2Þ Using Du ¼ 1  pu gives the formula in Table 1.
log ð1 xÞ
In addition,
Continuous log-series
X
1
1x
nψ ðnÞ ¼  ; ðA:3Þ
n¼1
xlog 1=x In this case, GðmÞ ¼ u is

X
1 ð1  xÞm  1 ¼ u; ðA:15Þ
ð1  xÞ⌈nu ⌉
nψ ðnÞ ¼  ; ðA:4Þ
n ¼ ⌈nu ⌉
xlog 1=x such that

and log ðuÞ

nu ¼ 1 þ ðA:16Þ
log ð1  xÞ
X
1
ð1 xÞ ⌈nu ⌉
ψ ðnÞ ¼   Φð1  x; 1; ⌈nu ⌉Þ: ðA:5Þ (this is the same as in the discrete case (Eq. (A.2)). Therefore,
n ¼ ⌈nu ⌉
log 1=x
Z 1  
li ð1  xÞnu liðuð1  xÞÞ
Thus, using Eq. (5), pu ¼ ψ ðnÞdn ¼ ¼ : ðA:17Þ
n ¼ nu lið1  xÞ lið1  xÞ
uð1  xÞ  ð1  xÞ⌈nu ⌉ ð1  xÞ⌈nu ⌉
pu ¼ þ Φð1  x; 1; ⌈nu ⌉Þ Using Du ¼ 1  pu gives the formula in Table 1.
xlog ð1=xÞð⌈nu ⌉  1Þ log ð1=xÞ
" #
ð1  xÞ u  ð1  xÞ⌈nu ⌉  1
¼ þ ð1  xÞ⌈nu ⌉  1 Φð1  x; 1; ⌈nu ⌉Þ : Continuous exponential
log ð1=xÞ xð⌈nu ⌉  1Þ
ðA:6Þ GðmÞ ¼ u becomes
Using Du ¼ 1  pu gives the formula for Du in Table 1. Further-  
eλð  m þ δÞ 1 þ λm
more, ¼ u: ðA:18Þ
1 þ δλ
X
1
ð1 xÞnu
pu ¼ ψ ðnÞ ¼  Φð1  x; 1; nu Þ Solving for m gives
n ¼ nu log 1=x

 
uð1 xÞ log ðuÞ  1  W  1  e  1  δλ uð1 þ δλÞ
¼  Φ 1  x; 1; 1 þ : ðA:7Þ nu ¼ ; ðA:19Þ
log 1=x log ð1  xÞ λ
Using Du ¼ 1  pu gives the formula for Du in Table 1. such that
Z 1 

uð1 þ δλÞ
Discrete exponential
pu ¼ ψ ðnÞdn ¼ eλð  nu þ δÞ ¼ exp 1 þ δλ þ W  1  ð1 þ δλÞ :
n ¼ nu e
FðmÞ ¼ u ðA:20Þ

is Using Du ¼ 1  pu gives the formula in Table 1.

   
e  λm 1 þ eλ  1 m ¼ u; ðA:8Þ Continuous gamma
such that
Here, GðmÞ ¼ u is equivalent to
1 W  1 ðuye  y Þ  
nu ¼  ; ðA:9Þ Γ 1 þ α; β m
1e λ λ ¼ u; ðA:21Þ
  Γð1 þ αÞ
where y ¼ λ= 1  eλ . In addition,
X
1
eλ which on rearranging gives
nψ ðnÞ ¼ λ ; ðA:10Þ  
n¼1
e 1 Γ 1 þ α; βm ¼ uΓð1 þ αÞ; ðA:22Þ
 T. Fung et al. / Journal of Theoretical Biology 386 (2015) 147–158 157

such that Continuous exponential

Q 1
ð1 þ α ; uÞ
nu ¼ : ðA:23Þ The dominance formula for the continuous exponential dis-
β tribution is:
Thus, h  i
  Du ¼ 1  exp 1 þ δλ þW  1  uð1 þ δλÞe  ð1 þ δλÞ : ðB:6Þ
Z  
Γ α; βnu Γ α; Q ð1 þ α; uÞ
1
1
pu ¼ ψ ðnÞdn ¼ ¼ : ðA:24Þ With δ 4 0,
n ¼ nu ΓðαÞ ΓðαÞ   
d 1 þ δλ þ W  1  uð1 þ δλÞe  ð1 þ δλÞ
Using Du ¼ 1  pu gives the formula in Table 1. dλ
(
  )
δλ W  1  uð1 þ δλÞe  ð1 þ δλÞ
Continuous lognormal ¼ δ 1   
1 þ δλ 1 þ W  1  uð1 þ δλÞe  ð1 þ δλÞ
(  )
In this case, GðmÞ ¼ u is δ 1 þ δλ þ W  1  uð1 þ δλÞe  ð1 þ δλÞ
¼   : ðB:7Þ

1 þ δλ 1 þ W  1  uð1 þ δλÞe  ð1 þ δλÞ
1 μ þ σ 2  log ðmÞ
1  erfc pffiffiffi ¼ u; ðA:25Þ
2 σ 2 The denominator of this derivative is thus negative, so that the
derivative has the opposite sign to 1 þ δλ þW  1 ð  uð1
such that
h i þ δλÞe  ð1 þ δλÞ Þ. Also, because δ 4 0, the derivative is non-zero and
pffiffiffi 1
nu ¼ exp μ þ σ 2  σ 2erfc ð2ð1  uÞÞ : ðA:26Þ therefore its sign does not change  with λ. For small
 positive λ,
1 þ δλ þ W  1 uð1 þ δλÞe  ð1 þ δλÞ  1 þ W  1  ue  1 o0 and the
Therefore, derivative is positive; hence the derivative is positive for all λ 4 0.
 
Z 1

This implies that dDu =dλ o 0, i.e. dDu =d J=S 40.

1 μ  log ðnu Þ
pu ¼ ψ ðnÞdn ¼ 2 erfc pffiffiffi
n ¼ nu 2 σ 2 Continuous lognormal

1 σ 1
¼ erfc pffiffiffi  erfc ð2ð1  uÞÞ : ðA:27Þ
2 2 Consider the dominance formula for the continuous lognormal
distribution:
Using Du ¼ 1  pu gives the formula in Table 1.

1 σ 1
Du ¼ 1  erfc pffiffiffi erfc ð2ð1  uÞÞ : ðB:8Þ
2 2
Appendix B. Details on mathematical analyses of dominance The complementary error function erfcðzÞ is a decreasing
 
formulae function of z, so dDu =dσ 4 0, i.e. dDu =d J=S 4 0.

Continuous log-series
Appendix C. Supplementary material
Consider the dominance formula for the continuous log-series
distribution: Supplementary data associated with this article can be found in
liðuð1  xÞÞ the online version at http://dx.doi.org/10.1016/j.jtbi.2015.09.011.
Du ¼ 1  : ðB:1Þ
lið1  xÞ

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