PERSPECTIVE

PUBLISHED: 22 JUNE 2017 | VOLUME: 1 | ARTICLE NUMBER: 0168

Revisiting the biodiversity–ecosystem

multifunctionality relationship
Lars Gamfeldt†* and Fabian Roger†*
A recent and prominent claim for the value of biodiversity is its importance for sustaining multiple ecosystem functions. The
general idea is intuitively appealing: since all species are to some extent unique, each will be important for a different set of
functions. Therefore, as more functions are considered, a greater diversity of species is necessary to sustain all functions
simultaneously. However, we show here that the relationship between biodiversity and ecosystem functioning does not change
with the number of functions considered. Biodiversity affects the level of multifunctionality via non-additive effects on indi-
vidual functions, and the effect on multifunctionality equals the average effect on single functions. These insights run counter
to messages in the literature. In the light of our simulations, we present limitations and pitfalls with current methods used to
study biodiversity–multifunctionality, which together provide a perspective for future studies.

B
iodiversity is causally linked to ecosystem functioning: changes
in the number of species in a community generally result in
changes in a range of functions1–5. Functioning increases with
species richness due to niche complementarity, positive interac-
tions, or the presence of certain influential species. The relationship
between richness and functioning can generally be described using
some version of a positive but decelerating function6,7, such that the
relationship is steep at low richness, and levels off quickly as richness
increases further.
Because species differ in their contributions to different func-
tions, it has been proposed that biodiversity should be more impor-
tant—and that the relationship should be less saturating—when
multiple functions are considered. This idea, often referred to as
multifunctionality 8, was originally proposed in a study on seagrass,
where it was observed that a diverse assemblage of small marine
grazers simultaneously maximized multiple functions9. Positive
biodiversity–multifunctionality relationships were later shown
for many different systems8,10–23, and a recent meta-analysis sug-
gests that a positive biodiversity–multifunctionality relationship
is general24.
How biodiversity influences ecosystem multifunctionality
Because all species are to some extent functionally unique, high
biodiversity may provide for a high variety of functions. If species
specialize in different functions, all species are needed to perform
all of the functions.
In addition to providing for more functions, increasing bio­
diversity is proposed to increase the level of multifunctionality.
More importantly, it is hypothesized that the importance of bio­
diversity increases with the number of functions considered. It is
this hypothesis that suggests that, as we increase the number of
functions that we study, the slope between biodiversity and multi­
functionality becomes steeper. The literature on biodiversity and
ecosystem multi­ functionality has mainly focused on this ‘bio­
diversity begets the level of multifunctionality’ hypothesis (for
example, refs 8,10,12,24–26), and, with few exceptions22,27,28, all papers
on the importance of biodiversity for multifunctionality study sets
of functions shared by all focal species. We should point out that
the case in which certain functions are unique to particular species
is a special case within the broader ‘biodiversity begets the level of
multifunctionality’ hypothesis. There is no conceptual difference
between a species that performs a given function at a very low level
and a species that does not perform the function at all. They are both
part of the same continuum. A value of zero will simply decrease the
average level of multifunctionality slightly more than a low value.
While the assumption that the slope between biodiversity and
multifunctionality becomes steeper with the number of functions is
not always mentioned explicitly, it is still an implicit assumption in
many papers. It is common for articles to include variations of state-
ments like “the impact of diversity is stronger when multiple func-
tions are considered together,” or “more species are needed to sustain
multiple functions than any single function,” or “studies focusing
on individual functions will underestimate levels of biodiversity
required for multifunctionality” (for example, refs 3,8,10,11,24,25,29,30).
The truth of such statements requires that the slope of the diversity–
ecosystem function relationship changes as we move from single to
multiple functions.
The importance of biodiversity for multifunctionality is
grounded on a verbal framework that makes intuitive sense. Species
have trade-offs in terms of allocating resources to growth, reproduc-
tion and survival, so no single species can maximize all ecosystem
functions9,31,32. A variety of organisms are hence required for an eco-
system to simultaneously sustain multiple functions. However, this
intuitive knowledge, blended with vague terminology, has painted a
hazy picture of a causal relationship between biodiversity and mul-
tifunctionality. We argue here that it is time scientists abandon the
idea that the effect on multifunctionality is larger than the average
effect on single functions, and that the importance of biodiversity is
dependent on the number of functions considered. We show that,
contrary to common belief, increasing the number of functions
considered does not by itself change the nature of the relationship
between biodiversity and ecosystem multifunctionality.
Defining biodiversity and multifunctionality
In order to assess the relationship between biodiversity and multi-
functionality, both variables first need to be defined. To make our
discussion easy to follow, we stick with species richness as our meas-
ure of diversity. Species richness is not a particularly good metric

Department of Marine Sciences, University of Gothenburg, Box 461, SE-405 30 Göteborg, Sweden. †These authors contributed equally to this work.
*e-mail: lars.gamfeldt@gu.se; fabian.roger@gu.se

NATURE ECOLOGY & EVOLUTION 1, 0168 (2017) | DOI: 10.1038/s41559-017-0168 | www.nature.com/natecolevol 1

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PERSPECTIVE NATURE ECOLOGY & EVOLUTION

Box 1 | Different ways of relating multifunctionality to changes in biodiversity.

To understand how biodiversity can be analytically related to

a b 30%
multi­functionality, it is useful to provide a hypothetical example F1
F2 3
(illustrated in the figure). Say that there are three species (X, Y, Z) 1.0

Number of functions ≥ threshold

F3 2
and three functions (F1, F2, F3), with each species providing the Average 1
three functions at different levels (green bars). We here set these
0
three function levels to be 0.2, 0.6 and 1.0. Species X is most

Function level
important for function 1, species Y for function 2, and species 0.6
Z for function 3. The average value for the three functions is the c 70%
same across all species (blue bars). Assuming no niches and equal 3
fitness across species (hence no biodiversity effects), the three- 2
species mixture XYZ has the same average function value as the 0.2 1
single species. If we are concerned with how many functions are 0
provided above a certain threshold of the maximum function lev- X Y Z XYZ 1 3
els, another picture emerges. At a level of 30% of the maximum, a Species composition Species richness
mean of two functions are above the threshold for single species
whereas all three functions are above the threshold for the three- Conceptual example with three species and functions. a, The figure
species mixture; a positive biodiversity effect. At a threshold level shows species X, Y and Z, which perform the functions F1, F2 and F3
of 70%, a mean of one function is above the threshold for single (green bars), and the average of those three functions (blue bars) for
species whereas no function is above the threshold for the mix- monocultures and the mixture. b,c, The number of functions sustained
ture; a negative relationship. This pattern has been described as the above a certain threshold (b, 30%; c, 70%) for monocultures and
jack-of-all-trades effect 23 (see main text). the mixture.

of diversity 33,34 and might not be appropriate under most circum-
stances, especially if diversity is observed and not manipulated. Yet
it is the simplest metric and will help us establish our framework,
which can be extended to other diversity metrics as desired.
There are at least four distinct approaches to analyse multi-
functionality: the species turnover approach8,13, the averaging
approach16,35, the threshold approach10,12,25, and the multivariate
modelling framework26. In this paper, we use the average and mul-
tiple threshold approaches to present the null expectation for bio-
diversity and multifunctionality. The averaging approach calculates
multifunctionality as the average of all standardized function val-
ues. The threshold approach tallies the number of functions that are
above one or multiple thresholds, usually defined as some percent-
age (often between 1% and 99%) of the maximum observed perfor-
mance for a given function. The two approaches give complementary
information, as explained in detail in Box 1.
The biodiversity–multifunctionality relationship
To explore the biodiversity multifunctionality–relationship, we sim-
ulate 15 species (A–O) that perform 9 functions (F1 to F9). The levels
at which each function is performed by each species is drawn from
a uniform distribution, U(0,1). Pairwise correlations between func-
tions vary randomly from negative (–0.5) to positive (0.6) with an
average correlation close to 0. The nine functions are standardized
by their maximum (f(x) = x / max(x)). In a second step we simulate a
diversity experiment where we combine the species from the species
pool in communities with richness 1 to 15. All possible communities
are simulated at each richness level. We use a replacement design,
that is, with species abundances declining proportionally with the
number of species. This means, for instance, that in a ten-species
mixture, each species constitutes 10% of the total abundance. This
is the most commonly used design in biodiversity ecosystem func-
tioning experiments and simulates a scenario with no niches and
equal fitness across species (under the assumption that all species
are equally abundant). Such a scenario yields the null expectation
used in biodiversity–ecosystem functioning studies when testing for
biodiversity effects. For example, to calculate selection and comple-
mentarity (sensu ref. 36), the proportion of each species in the species
mixture, assuming equal abundance, is used to calculate expected
values of functions, based on species performances in monoculture.
We consider two scenarios. First we consider a scenario in which
each individual function is unrelated to biodiversity (Fig. 1a). For
any level of richness, the average function value is the average of
all possible species combinations, which is simply the average func-
tion value across all species in the species pool. The mean function
values for all single functions thus stay the same regardless of spe-
cies richness. Since average multifunctionality is calculated as the
average of all standardized single functions, it too does not change
with diversity (Fig. 1b). Considering the functions jointly yields the
average of all nine functions.
If we are interested in the number of functions above a certain
threshold level (expressed as the percentage of the maximum level
of that function across all species combinations and richness lev-
els) the expectation depends on the threshold level. The diversity
effect is positive at low thresholds (below roughly the grand mean of
all functions, here 52%), but at thresholds above that, the diversity
effect is negative (Fig. 1c). The patterns in Fig. 1c mirror those in
Box 1. An alternative way of illustrating the patterns in Fig. 1c is to
plot the slope of the relationship between species richness and the
number of functions above the full range of thresholds25 (Fig. 1d).
Point p in Fig. 1d shows that the slope is zero at the 51% threshold.
With the threshold approach, we observe that biodiversity can be
both positively and negatively related to multifunctionality without
any non-additive interactions. Non-additive effects include positive
and negative species interactions, which alone or in combination
result in what is referred to as biodiversity effects36. For a scenario
with no niches and equal fitness across species, there is a positive
relationship at thresholds below roughly the grand mean across all
functions and a negative relationship above that. The steepness of
the slope depends on the threshold level (Fig. 1b).
The results presented in Fig. 1a–d are similar to those provided in a
recent paper on forest biodiversity and ecosystem multi­functionality 23.
The authors propose that these patterns are driven by a mechanism
referred to as the jack-of-all-trades effect. The English saying ‘jack of
all trades, master of none’ is taken as metaphor for the description
of the pattern that diverse communities can uphold many functions
above low threshold levels, but only few functions above high thresh-
old levels. Species effects on different functions average out in species
mixtures, so that communities with many species will, according to
the threshold approach, have higher multifunctionality for low levels

2 NATURE ECOLOGY & EVOLUTION 1, 0168 (2017) | DOI: 10.1038/s41559-017-0168 | www.nature.com/natecolevol

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NATURE ECOLOGY & EVOLUTION PERSPECTIVE
a e
F1 F2 F3 F1 F2 F3
1.0 1.0
0.5 0.5
0.0
Value of function

Value of function
0.0
F4 F5 F6 F4 F5 F6
1.0 1.0
0.5 0.5
0.0 0.0
F7 F8 F9 F7 F8 F9
1.0 1.0
0.5 0.5
0.0 0.0
4 8 12 4 8 12 4 8 12 4 8 12 4 8 12 4 8 12
Species richness Species richness
b f
Average value of standardized functions

Average value of standardized functions

0.7

0.6
0.6

0.5
0.5

0.4
0.4
1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15
Species richness Species richness
c 10% 20% 30%
g 10% 20% 30%
9 9
6 6
Number of functions ≥ threshold

Number of functions ≥ threshold

3 3
0 0
40% 50% 60% 40% 50% 60%
9 9
6 6
3 3
0 0
70% 80% 90% 70% 80% 90%
9 9
6 6
3 3
0 0
1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15
Species richness Species richness
d h

0.2 0.2
p: 52% p: 62%
Slope
Slope

0.0 0.0

-0.2 –0.2

25 50 75 100 25 50 75 100
Threshold (%) Threshold (%)

Figure 1 | Two scenarios for the relationship between biodiversity and multifunctionality. a–h, The relationship between biodiversity (species richness)
and multifunctionality for a scenario with no species interactions (a-d) and with complementarity for two functions (1 and 6, e–h). Relationships are shown
for individual functions (a,e), average function level (b,f) and for the number of functions above or equal to each of nine thresholds (c,g). Also presented
is how the slope of the relationship between biodiversity and number of functions above threshold changes with threshold level (d,h). In plots a–c and
e–g, point values are rounded to the third decimal and only unique points are shown. Lighter colours represent higher point densities. The regression line
is fitted to all points before rounding (n = 32,767). In plots d and h, the shaded area represents the 95% confidence interval of the slope estimates, point p
indicates the threshold at which the slope turns from positive to negative.

of functioning, but lower multifunctionality if high levels of function- diversity due to the statistical averaging of fluctuations in species’
ing are required. The jack-of-all-trades effect is explained as broadly abundances. While the simulations in the forest multifunctionality
analogous to the averaging effect underlying the portfolio effect 23. The paper 23 and our study reveal similar patterns, we offer a very different
portfolio effect (sensu ref. 37) means that stability rises with species interpretation as we discuss in more detail later.

NATURE ECOLOGY & EVOLUTION 1, 0168 (2017) | DOI: 10.1038/s41559-017-0168 | www.nature.com/natecolevol 3

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PERSPECTIVE NATURE ECOLOGY & EVOLUTION

a b c

0.03 0.25 0.25

Slope estimate

Slope estimate
Slope estimate
0.02
0.00 0.00

0.01
–0.25 –0.25

0.00
1 2 3 4 5 6 7 8 9 25 50 75 100 25 50 75 100
Number of functions considered Threshold (%) Threshold (%)
Number of functions 0 3 Number of functions 3 6 Number of species 6 9
with complementarity 6 9 (15 species) 9 (9 functions) 15

Figure 2 | The slope between species richness and average multifunctionality in relation to the number of functions and species considered. a, The
figure shows the slopes for the average multifunctionality–diversity relationship (for 1–12 species) for all possible combinations of 9 functions. We present
4 scenarios in which there is complementarity for 0, 3, 6 and 9 functions (out of 9 total). The average slope increases with the number of functions subject
to complementarity, but does not change with the number of functions included. Although individual slopes vary depending on exactly which functions
are included in any one combination of functions, the average slope stays constant over the full range of number of functions. For each scenario and each
number of functions n we plot the slope for all possible combinations of n out of 9 functions, 511 slopes in total per scenario. b, Changes in the effect
of biodiversity as a function of the number of functions using the multi-threshold approach. Curves for 3, 6 and 9 functions are shown holding species
richness constant at 15 species. Maximum and minimum slopes increase as more functions are included. Lines are means ± 95% confidence interval of the
slopes calculated for all possible combinations of n (= 3, 6 or 9) out of 9 functions. c, An equivalent plot to that in b but for three different richness levels,
holding the number of functions constant. It shows that the maximum and minimum slopes decrease with increasing maximum species richness. Lines are
means ± 95% confidence interval of the slopes calculated for all possible combinations of n (= 6, 9 or 15) out of 15 species.

In the second scenario that we simulate, we include a positive
complementarity effect. In our example, we assign complementa-
rity to two of the nine functions (functions 1 and 6, Fig. 1e). We
could have chosen an example including any number of functions
subject to complementarity, since our approach is general (as we
show in ‘Number of functions does not matter’ below). We imple-
ment complementarity by multiplying the single function values by
a complementarity factor (CF), where the complementarity factor is
a saturating function of species richness (S). The complementarity
factor is 1 for monocultures and grows to a predefined maximum
(CFmax), which was set to 3 in our example.
CF = CFmax × (1 – (1 – 1 / CFmax) × exp(1 – Sr))
where r defines the speed at which CF grows towards CFmax and is
set to 0.5.
Complementarity causes the average multifunctionality to be
positively related to species richness (Fig. 1f). The slope of the aver-
age multifunctionality relationship is the average of the seven flat
and the two positive individual function slopes. The complemen-
tarity for two functions also causes the upper range of the positive
multiple threshold effect to be pushed towards higher threshold lev-
els (here around 63%, see Fig. 1g,h). In other words, we observe a
positive effect of biodiversity over a wider range of thresholds. A
negative effect is observed for thresholds above 68%.
Number of functions does not matter
The scenario in which biodiversity is unrelated to each individual
function demonstrates that biodiversity does not increase the aver-
age level of multiple functions (Fig. 1b). Considering more func-
tions does not affect the relationship between biodiversity and
multifunctionality, as it is often claimed. A direct way of showing
this is by plotting the slope between species richness and average
multifunctionality on the y-axis against the number of functions
on the x-axis. Figure 2a shows the slopes for the scenario in which
biodiversity is unrelated to each individual function, as well as for
scenarios in which there is complementarity for three, six and nine
functions (compare with Fig. 1e–h for a scenario with two func-
tions with complementarity). As expected, the average slope of the
diversity–multifunctionality relationship increases with the number
of functions subject to complementarity. However, the average slope
does not change with the total number of functions considered.
Although individual slopes vary depending on exactly which func-
tions are included in any one combination of functions, the average
slope stays constant across the full range of number of functions.
What does it look like if we instead use the threshold approach
to study how the effect of biodiversity changes with the number of
functions? Figure 2b shows the slope of the relationship between
the number of functions above thresholds and species richness
against the threshold (as in Fig. 1d,h). We present curves for three,
six and nine functions, holding species richness constant at 15.
Maximum and minimum slopes increase as more functions are
included. Figure 2c, which presents a similar plot but for three dif-
ferent richness levels, holding the number of functions constant,
shows that the maximum and minimum slopes decrease with
increasing richness. While the patterns in Fig. 2 seem intriguing,
they are simply due to the fact that we vary the range on the y-axis
while holding the x-axis constant (panel b—an increasing number
of functions with constant richness) or vary the range on the x-axis
while holding the y-axis constant (panel c—an increasing number
of species with constant number of functions). A change in the
maximum effect of diversity does thus not represent any under­lying
biological phenomenon. One way to illustrate this point is to con-
sider a hypothetical example with ten species and nine functions
(Supplementary Fig. 1). All functions are perfectly and positively
correlated, meaning that for each species the contribution is the
same for all functions. This is equivalent to studying a single func-
tion. Regardless of how many of the nine functions we subtract or
add, we still effectively study a single function. Yet, we observe a
change in slopes, depending on how many functions we include.
Furthermore, if Fig. 2b should be taken as evidence for stronger
diversity effects at high number of functions we must conclude, by
the same logic, that considering more species decreases the effect of
biodiversity on multifunctionality.

4 NATURE ECOLOGY & EVOLUTION 1, 0168 (2017) | DOI: 10.1038/s41559-017-0168 | www.nature.com/natecolevol

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NATURE ECOLOGY & EVOLUTION PERSPECTIVE
a
0.4

0.2
Slope estimate

0.0

Scaled by max
–0.2 Scaled (0,1)

25 50 75 100
Threshold (%)

b c
4 f(x) = x – min(x) 6 f(x) = x
max(x) – min(x) max(x)
3
4

Density
Density

2
1

0 0
0.00 0.25 0.50 0.75 1.00 0.00 0.25 0.50 0.75 1.00
Scaled between (0,1) Scaled by maximum

Figure 3 | The effect of standardization choice on the result of the multi-threshold approach. a, The expected result from the multi-threshold approach
for the German site of the BIODEPTH project using two different methods to standardize raw function values. We used real data from the monocultures
to simulate the null expectation for the full richness gradient. We standardized the data to be between 0 and 1 (blue line) or by the maximum (red line).
Both standardization techniques are common in the literature but the choice has a large impact on the resulting slope pattern. The shaded areas represent
the 95% confidence interval of the slope estimates (n = 1,023); the simulation was fully replicated (all possible species composition at each richness
level). b and c show the effect of the two different standardization techniques on the mean and the distribution of the functions, which results in the two
distinct patterns. The dashed lines in b and c mark the grand mean of all standardized function values.

The devil is in the multifunctionality detail
The fact that the relationship between biodiversity and multi­
functionality is sensitive to the number of functions and species,
makes it clear that we need to be careful when interpreting the
results of any one study. But these are not the only factors that com-
plicate interpretations. Because, as the saying goes: the devil is in the
detail. For example, the way we prepare our data for analyses will
have implications for the results. We can use the raw data, standard-
ize data by dividing by their maximum (or some top percentage of
the maximum), standardize them to range from 0 to 1, or convert
data to standard scores. If we compare two ways of standardization
we see that standardizing by the maximum value (f(x) = x / max(x))
produces a different pattern than if we standardize between 0 and 1
(f(x) = (x – min(x)) / (max(x) – min(x))). For the Germany site in
the BIODEPTH project 8,38,39, the expected slopes for the standardi-
zation relative to the maximum are generally less steep compared
to slopes based on the standardization between 0 and 1 (Fig. 3a).
The threshold at which the slope turns from positive to negative
(compare with point p in Fig. 1d,h) occurs at a higher threshold for
the standardization relative to the maximum (around 50%) com-
pared to the standardization between 0 and 1 (around 35%). This
is because the standardization of each function between 0 and 1
decreases the grand mean of all function values. The implications
of such apparently trivial choices have hitherto been overlooked in
the field. Distributions for the standardized data for the BIODEPTH
Germany site are shown in Fig. 3b (standardized between 0 and 1)
and Fig. 3c (standardized to the maximum). Raw data are available
via the multifunc package25.
While we chose to make the point above using experimental
data from BIODEPTH, other examples can easily be simulated. If,
for example, function values are drawn from a normal distribution
such that no function value is close to 0 (for example, mean = 30,
standard deviation = 6), the standardization by the maximum (or
for that matter the equivalent choice of using unstandardized data)
will show a dramatically different pattern compared to a standardi-
zation between 0 and 1. The latter will show the now familiar pat-
tern of positive slopes below 50% and negative slopes above. The
former, however, yields flat slopes (that is, no effect of biodiversity)
up to a threshold of 50%, followed by the typical pattern of positive
(50–76%) and negative slopes (>76%, Supplementary Fig. 2). Note
that in this hypothetical example, the range of threshold values with
the highest positive slopes for the standardization by the maximum
is the range of threshold values with the highest negative slopes for
the standardization between 0 and 1. We would like to add that
standardizing functions between 0 and 1 has strong implications for
the interpretation: if we take the example of a function with values
ranging from 90 to 100, then, by standardizing it between 0 and 1, we
effectively say that the species that performs the function at level 90,
does not perform the function at all and that the species performing
the function at level 95 performs it at 50% of its maximum.
It is clear that the results from any one study on multifunctional-
ity will be sensitive to which exact approach is used. An important
message emerging from our simulations is hence that studies on
multifunctionality need to be explicit about their methods and null
expectations (for example, given the performance of monoculture).
Such null expectations are currently absent in the multifunctionality
field. Furthermore, not only will the expected distributions depend
on standardization, but also on the original distributions of each
individual function, as well as on how many functions and species
are included (Fig. 2). Since the slope of the relationship is sensitive
to the number of functions and species, caution should be taken
when comparing biodiversity effects across studies. The full range

NATURE ECOLOGY & EVOLUTION 1, 0168 (2017) | DOI: 10.1038/s41559-017-0168 | www.nature.com/natecolevol 5

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PERSPECTIVE NATURE ECOLOGY & EVOLUTION

of null-expectations observed during our simulations goes beyond
what we have space to discuss here. They make it clear, however, that
the null expectations for the multi-threshold approach vary widely
and are non-intuitive. We therefore caution against the use of the
multi-threshold approach as our simulations indicate that a coher-
ent interpretation is challenging. As this conclusion does not invali-
date the justified criticism of alternative multifunctionality metrics
articulated previously 25, we think it is time for the field of multi-
functionality to reconsider its current use of metrics. We provide an
online simulation tool where a multitude of different scenarios can
be simulated. It is our hope that the online application can aid the
understanding of current and future metrics of multifunctionality
(see ‘Code availability’).
Concluding remarks
Even though species differ in their importance for different func-
tions, we demonstrate that the effect on multifunctionality is not
larger than the average effect on single functions. Quite the con-
trary, it is sobering to realize that in the absence of positive diver-
sity effects, multifunctionality can almost always be maximized by
the highest performing monoculture, just as with single functions
(Fig. 1b,c). Regressions on the 100% quantile of our simulated data
reveal that the relationship between biodiversity and multifunction-
ality goes from flat at low thresholds to negative at high thresholds
(Supplementary Fig. 3), and is never positive.
As we have demonstrated, claims that considering multiple func-
tions increases the need for, and therefore the value of, biodiversity
are not justified. Many terms in the biodiversity–multifunctionality
literature give fuel to such claims. Examples of terms include “com-
plementarity across multiple functions,”10 “multivariate dominance
effect,”9 and “multivariate complementarity”40. We emphasize that
these, and similar, terms are misleading. There can only be selec-
tion and complementarity for single functions, and mechanisms
behind high levels of multifunctionality must thus also be sought
in single functions.
The natural world is, of course, often not characterized by equal
abundances of all species and the absence of niches and fitness dif-
ferences. Increases in biodiversity, for example, by the addition or
arrival of new species to an ecosystem, often does not translate to
a direct and proportional decline in the already present species41
(which is the case in a replacement design). And conversely, loss of
species may not result in a compensatory and proportional increase
in the remaining species. In such cases, what we have is a bio­
diversity effect in terms of niche complementarity for abundance
and/or biomass production. Such complementarity has strong
potential to drive changes in other functions. For example, higher
abundance and/or biomass of organisms within and across trophic
levels can drive increases in fluxes of material and energy 42,43.
If we are to better manage ecosystems and the services they pro-
vide we need a thorough understanding of the role that biodiversity
plays. To quote Cardinale et al.44: “the three most important fac-
tors that will determine the success of the biodiversity–ecosystem
functioning paradigm will be our ability to identify mechanisms,
mechanisms, mechanisms!” We argue that this is very true for the
biodiversity–multifunctionality field. In that light, we need to learn
more about which functions are driven by biodiversity and how
they in turn affect other functions and multifunctionality. We are
not excluding the possibility that there might be a yet undiscovered
mechanism which directly links diversity and multifunctionality.
However, the burden of proof lies on us ecologists, and we must
stop assuming a mechanism where none has been shown.
We emphasize that we do not send the message that multi­
functionality should not be studied. It is indeed important to exam-
ine multiple aspects of functioning associated with biodiversity loss,
climate change, land use intensification, and other dimensions of
environmental change45–47. Multifunctionality metrics can be useful
in summarizing the collective behaviour of multiple ecosystem
functions and services, and in dealing with trade-offs25. It is quite
plausible, that when there is a fixed set of functions of interest,
with threshold levels that can be specified a priori, and the species
pool is defined, a threshold approach can be used to examine if and
when biodiversity is important to sustain multifunctionality. There
may, for example, be applied contexts in which we know that only
a defined small reduction is acceptable for production, whereas we
accept much larger decreases in cultural services. However, it is nec-
essary that we are aware that the importance of biodiversity does
not change as we move from single to multiple functions. Hand-
waving claims that the value of biodiversity increases with higher
dimensions of functioning will only lead research and management
efforts astray.
Code availability
The application can be accessed at: https://fabianroger.shinyapps.
io/multifunctionality-simulations/. The source code for the app
can be found here: https://github.com/FabianRoger/Gamfeldt_
Roger_2017_NEE_app and https://doi.org/10.5281/zenodo.496151.
The R code producing all simulations presented in this article
can be found here: https://github.com/FabianRoger/Gamfeldt_
Roger_2017_NEE and https://doi.org/10.5281/zenodo.496144. To
run from within R, execute:
library(shiny)
runGitHub(“FabianRoger/Gamfeldt_Roger_2017_NEE_app”)
Received 25 November 2016; accepted 19 April 2017;
published 22 June 2017
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Acknowledgements
We would like to thank M. Jonsson, J. Griffin and J. Bengtsson for providing valuable
feedback and comments on earlier drafts. This work was supported by grant 621-2009-
5457 from the Swedish research council VR to L.G.
Author contributions
Study idea by L.G.; L.G. and F.R. designed the study; F.R. wrote the script for the
analyses; L.G. and F.R. interpreted the analyses and wrote the paper.
Additional information
Supplementary information is available for this paper.
Reprints and permissions information is available at www.nature.com/reprints.
Correspondence should be addressed to L.G. and F.R.
How to cite this article: Gamfeldt, L. & Roger, F. Revisiting the biodiversity–ecosystem
multifunctionality relationship. Nat. Ecol. Evol. 1, 0168 (2017).
Publisher’s note: Springer Nature remains neutral with regard to jurisdictional claims in
published maps and institutional affiliations.
Competing interests
The authors declare no competing financial interests.

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