This document contains answers to some of the most frequently asked questions about R package vegan.
This work is licensed under the Creative Commons Attribution 3.0 License. To view a copy of this license, visit https://creativecommons.org/licenses/by/3.0/ or send a letter to Creative Commons, 543 Howard Street, 5th Floor, San Francisco, California, 94105, USA.
Copyright © 2008-2020 vegan development team
Vegan is an R package for community ecologists. It contains the most popular methods of multivariate analysis needed in analysing ecological communities, and tools for diversity analysis, and other potentially useful functions. Vegan is not self-contained but it must be run under R statistical environment, and it also depends on many other R packages. Vegan is free software and distributed under GPL2 license.
R is a system for statistical computation and graphics. It consists of a language plus a run-time environment with graphics, a debugger, access to certain system functions, and the ability to run programs stored in script files.
R has a home page at https://www.R-project.org/. It is free software distributed under a GNU-style copyleft, and an official part of the GNU project (“GNU S”).
Both R and latest release version of vegan can be obtained through CRAN. Unstable development version of vegan can be obtained through GitHub. Formerly vegan was developed in R-Forge, but after moving to GitHub the R-Forge repository is out of date.
Vegan depends on the permute package which will provide advanced and flexible permutation routines for vegan. The permute package is developed together with vegan in GitHub.
Some individual vegan functions depend on packages MASS, mgcv, parallel, cluster, lattice and tcltk. These all are base or recommended R packages that should be available in every R installation. Vegan declares these as suggested or imported packages, and you can install vegan and use most of its functions without these packages.
Vegan is accompanied with a supporting package vegan3d for three-dimensional and dynamic plotting. The vegan3d package needs non-standard packages rgl and scatterplot3d.
CRAN Task Views include
describe several useful packages and functions. If you install R package
ctv, you can inspect Task Views from your R session, and
automatically install sets of most important packages.
Vegan is a fully documented R package with standard help pages.
These are the most authoritative sources of documentation (and as a
last resource you can use the force and the read the source, as
vegan is open source). Vegan package ships with other
documents which can be read with
command. The documents included in the vegan package are
Web documents outside the package include:
Roeland Kindt has made package BiodiversityR which provides a GUI
for vegan. The package is available at
It is not a mere GUI for vegan, but adds some new functions and
complements vegan functions in order to provide a workbench for
biodiversity analysis. You can install BiodiversityR using
install.packages("BiodiversityR") or graphical package management menu
in R. The GUI works on Windows, MacOS X and Linux.
citation("vegan") in R to see the recommended citation to
be used in publications.
In general, you do not need to build vegan from sources, but binary builds of release versions are available through CRAN for Windows and MacOS X. If you use some other operating systems, you may have to use source packages. Vegan is a standard R package, and can be built like instructed in R documentation. Vegan contains source files in C and FORTRAN, and you need appropriate compilers (which may need more work in Windows and MacOS X).
Not currently. You need tools to build C and Fortran programs to
install vegan. If you have those, you can use
devtools::install_github("vegan") to install the most recent devel
If you think you have found a bug in vegan, you should report it to vegan maintainers or developers. The preferred forum to report bugs is GitHub. The bug report should be so detailed that the bug can be replicated and corrected. Preferably, you should send an example that causes a bug. If it needs a data set that is not available in R, you should send a minimal data set as well. You also should paste the output or error message in your message. You also should specify which version of vegan you used.
Bug reports are welcome: they are the only way to make vegan non-buggy.
Please note that you shall not send bug reports to R mailing lists, since vegan is not a standard R package.
It is not necessarily a bug if some function gives different results
than you expect: That may be a deliberate design decision. It may be
useful to check the documentation of the function to see what was the
intended behaviour. It may also happen that function has an argument to
switch the behaviour to match your expectation. For instance, function
vegdist always calculates quantitative indices (when this is
possible). If you expect it to calculate a binary index, you should use
binary = TRUE.
Vegan is dependent on user contribution. All feedback is welcome. If you have problems with vegan, it may be as simple as incomplete documentation, and we shall do our best to improve the documents.
Feature requests also are welcome, but they are not necessarily fulfilled. A new feature will be added if it is easy to do and it looks useful, or if you submit code.
If you can write code yourself, the best forum to contribute to vegan is GitHub.
You are wrong! Computers are painfully pedantic, and if they find
non-numeric or negative data entries, you really have them. Check your
data. Most common reasons for non-numeric data are that row names were
read as a non-numeric variable instead of being used as row names (check
row.names in reading the data), or that the column names were
interpreted as data (check argument
header = TRUE in reading the
data). Another common reason is that you had empty cells in your input
data, and these were interpreted as missing values.
Yes. Most vegan methods can handle binary data or cover abundance data. Most statistical tests are based on permutation, and do not make distributional assumptions. There are some methods (mainly in diversity analysis) that need count data. These methods check that input data are integers, but they may be fooled by cover class data.
Most commonly the reason is that other software use presence–absence
data whereas vegan used quantitative data. Usually vegan indices
are quantitative, but you can use argument
binary = TRUE to make them
presence–absence. However, the index name is the same in both cases,
although different names usually occur in literature. For instance,
Jaccard index actually refers to the binary index, but vegan uses
"jaccard" for the quantitative index, too.
Another reason may be that indices indeed are defined differently, because people use same names for different indices.
Stress is a proportional measure of badness of fit. The proportions can
be expressed either as parts of one or as percents. Function
(MASS package) uses percents, and function
package) uses proportions, and therefore the same stress is 100 times
isoMDS. The results of
goodness function also depend on
the definition of stress, and the same
goodness is 100 times higher in
isoMDS than in
monoMDS. Both of these conventions are equally
Most common reason is that you have too few observations for your NMDS.
n observations (points) and
k dimensions you need to estimate
n*k parameters (ordination scores) using
k dimensions you must have
n > 2*k + 1, or for two dimensions at
least six points. In some degenerate situations you may need even a
larger number of points. If you have a lower number of points, you can
find an undefined number of perfect (stress is zero) but different
solutions. Conventional wisdom due to Kruskal is that you should have
n > 4*k + 1 points for
k dimensions. A typical symptom of
insufficient data is that you have (nearly) zero stress but no two
convergent solutions. In those cases you should reduce the number of
k) and with very small data sets you should not use
NMDS, but rely on metric methods.
It seems that local and hybrid scaling with
monoMDS have similar lower
limits in practice (although theoretically they could differ). However,
higher number of dimensions can be used in metric scaling, both with
monoMDS and in principal coordinates analysis (
wcmdscale in vegan).
metaMDS uses function
monoMDS as its default method for
NMDS, and this function can handle zero dissimilarities. Alternative
isoMDS cannot handle zero dissimilarities. If you want to use
isoMDS, you can use argument
zerodist = "add" in
metaMDS to handle
zero dissimilarities. With this argument, zero dissimilarities are
replaced with a small positive value, and they can be handled in
isoMDS. This is a kluge, and some people do not like this. A more
principal solution is to remove duplicate sites using R command
unique. However, after some standardizations or with some
dissimilarity indices, originally non-unique sites can have zero
dissimilarity, and you have to resort to the kluge (or work harder with
your data). Usually it is better to use
Claims like this have indeed been at large in the Internet, but they are
based on grave misunderstanding and are plainly wrong. NMDS ordination
results are strictly metric, and in vegan
they are even strictly Euclidean. The method is called “non-metric”
because the Euclidean distances in ordination space have a non-metric
rank-order relationship to community dissimilarities. You can inspect
this non-linear step curve using function
stressplot in vegan.
Because the ordination scores are strictly Euclidean, it is correct to
use vegan functions
ordisurf with NMDS results.
Normally you can use function
scores to extract ordination scores for
any ordination method. The
scores function can also find ordination
scores for many non-vegan functions such as for
princomp and for some ade4 functions.
In some cases the ordination result object stores raw scores, and the
axes are also scaled appropriate when you access them with
rda the ordination object has only so-called
normalized scores, and they are scaled for ordination plots or for other
use when they are accessed with
The scaling or RDA results indeed differ from most other software
packages. The scaling of RDA is such a complicated issue that it cannot
be explained in this FAQ, but it is explained in a separate pdf document
on “Design decision and implementation details in vegan” that you can
read with command
If the RDA ordination results have a weird format or you cannot plot
them properly, you probably have a name clash with klaR package
which also has function
rda, and the klaR
predict functions are used for vegan RDA results. You can choose
rda functions using
will get obscure error messages if you use the wrong function. In
general, vegan should be able to work normally if vegan was
loaded after klaR, but if klaR was loaded later, its functions
will take precedence over vegan. Sometimes vegan namespace is
loaded automatically when restoring a previously stored workspace at
the start-up, and then klaR methods will always take precedence
over vegan. You should check your loaded packages. klaR may be
also loaded indirectly via other packages (in the reported cases it
was most often loaded via agricolae package). Vegan and
klaR both have the same function name (
rda), and it may not be
possible to use these packages simultaneously, and the safest choice
is to unload one of the packages if only possible. See discussion in
vegan github issues.
If you have a very old version of ade4 (prior to 1.7-8), you may
have similar name clashes with
cca. The solution is to upgrade
Constrained ordination (
capscale) will sometimes fail
with error message
Error in La.svd(x, nu, nv): error code 1 from Lapack routine 'dgesdd'.
It seems that the basic problem is in the
svd function of
that is used for numerical analysis in R.
LAPACK is an external
library that is beyond the control of package developers and R core team
so that these problems may be unsolvable. It seems that the problems
LAPACK code are so common that even the help page of
warns about them
Reducing the range of constraints (environmental variables) helps
sometimes. For instance, multiplying constraints by a constant < 1.
This rescaling does not influence the numerical results of constrained
ordination, but it can complicate further analyses when values of
constraints are needed, because the same scaling must be applied there.
We can only hope that this problem is fixed in the future versions of R
In general, vegan does not directly give any statistics on the “variance explained” by ordination axes or by the constrained axes. This is a design decision: I think this information is normally useless and often misleading. In community ordination, the goal typically is not to explain the variance, but to find the “gradients” or main trends in the data. The “total variation” often is meaningless, and all proportions of meaningless values also are meaningless. Often a better solution explains a smaller part of “total variation”. For instance, in unstandardized principal components analysis most of the variance is generated by a small number of most abundant species, and they are easy to “explain” because data really are not very multivariate. If you standardize your data, all species are equally important. The first axes explains much less of the “total variation”, but now they explain all species equally, and results typically are much more useful for the whole community. Correspondence analysis uses another measure of variation (which is not variance), and again it typically explains a “smaller proportion” than principal components but with a better result. Detrended correspondence analysis and nonmetric multidimensional scaling even do not try to “explain” the variation, but use other criteria. All methods are incommensurable, and it is impossible to compare methods using “explanation of variation”.
If you still want to get “explanation of variation” (or a deranged editor requests that from you), it is possible to get this information for some methods:
capscalegive the variation of conditional (partialled), constrained (canonical) and residual components, but you must calculate the proportions by hand. Function
eigenvalsextracts the eigenvalues, and
summary(eigenvals(ord))reports the proportions explained in the result object
RsquareAdjgives the R-squared and adjusted R-squared (if available) for constrained components. Function
goodnessgives the same statistics for individual species or sites (species are unavailable with
capscale). In addition, there is a special function
varpartfor unbiased partitioning of variance between up to four separate components in redundancy analysis.
decorana). The total amount of variation is undefined in detrended correspondence analysis, and therefore proportions from total are unknown and undefined. DCA is not a method for decomposition of variation, and therefore these proportions would not make sense either.
stressplotdisplays the nonlinear fit and gives this statistic.
No. Strictly speaking, this is impossible. However, you can define models that respond to similar goals as random effects models, although they strictly speaking use only fixed effects.
Constrained ordination functions
capscale can have
Condition() terms in their formula. The
Condition() define partial
terms that are fitted before other constraints and can be used to remove
the effects of background variables, and their contribution to
decomposing inertia (variance) is reported separately. These partial
terms are often regarded as similar to random effects, but they are
still fitted in the same way as other terms and strictly speaking they
are fixed terms.
adonis evaluates terms sequentially. In a model with
~ A + B the effects of
A are evaluated first, and
the effects of
B after removing the effects of
A. Sequential tests
are also available in
anova function for constrained ordination
results by setting argument
by = "term". In this way, the first terms
can serve in a similar role as random effects, although they are fitted
in the same way as all other terms, and strictly speaking they are fixed
All permutation tests in vegan are based on the permute package
that allows constructing various restricted permutation schemes. For
instance, you can set levels of
blocks for a factor
regarded as a random term.
A major reason why real random effects models are impossible in most vegan functions is that their tests are based on the permutation of the data. The data are given, that is fixed, and therefore permutation tests are basically tests of fixed terms on fixed data. Random effect terms would require permutations of data with a random component instead of the given, fixed data, and such tests are not available in vegan.
Vegan does not have a concept of passive points, or a point that
should only little influence the ordination results. However, you can
add points to eigenvector methods using
predict functions with
newdata. You can first perform an ordination without some species or
sites, and then you can find scores for all points using your complete
predict functions are available for basic
eigenvector methods in vegan (
decorana, for an
up-to-date list, use command
methods("predict")). You also can
simulate the passive points in R by using low weights to row and columns
(this is the method used in software with passive points). For instance,
the following command makes row 3 “passive”:
dune[3,] <- 0.001*dune[3,].
You should define a class variable as an R
factor, and vegan will
automatically handle them with formula interface. You also can define
constrained ordination without formula interface, but then you must code
your class variables by hand.
R (and vegan) knows both unordered and ordered factors. Unordered factors are internally coded as dummy variables, but one redundant level is removed or aliased. With default contrasts, the removed level is the first one. Ordered factors are expressed as polynomial contrasts. Both of these contrasts explained in standard R documentation.
The printed output of
envfit gives the direction cosines which are the
coordinates of unit length arrows. For plotting, these are scaled by
their correlation (square roots of column
r2). You can see the scaled
envfit arrows using command
The scaled environmental vectors from
envfit and the arrows for
continuous environmental variables in constrained ordination (
capscale) are adjusted to fill the current graph. The lengths
of arrows do not have fixed meaning with respect to the points (species,
sites), but they can only compared against each other, and therefore
only their relative lengths are important.
If you want change the scaling of the arrows, you can use
(plotting arrows and text) or
points (plotting only arrows) functions
for constrained ordination. These functions have argument
which sets the multiplier. The
plot function for
envfit also has the
arrow.mul argument to set the arrow multiplier. If you save the
invisible result of the constrained ordination
plot command, you can
see the value of the currently used
arrow.mul which is saved as an
ordiArrowMul is used to find the scaling for the current
plot. You can use this function to see how arrows would be scaled:
sol <- cca(varespec) ef <- envfit(sol ~ ., varechem) plot(sol) ordiArrowMul(scores(ef, display="vectors"))
vegan uses standard R utilities for defining contrasts. The default in
standard installations is to use treatment contrasts, but you can change
the behaviour globally setting
options or locally by using keyword
contrasts. Please check the R help pages and user manuals for details.
Aliased variable has no information because it can be expressed with the help of other variables. Such variables are automatically removed in constrained ordination in vegan. The aliased variables can be redundant levels of factors or whole variables.
alias gives the defining equations for aliased
variables. If you only want to see the names of aliased variables or
levels in solution
You can fit vectors or class centroids for aliased variables using
envfit function. The
envfit function uses weighted fitting, and the
fitted vectors are identical to the vectors in correspondence analysis.
Vegan uses permute package in all its permutation tests. The
permute package will allow restricted permutation designs for time
series, line transects, spatial grids and blocking factors. The
construction of restricted permutation schemes is explained in the
permutations in vegan and in the documentation of the
The default ordination
plot function is intended for fast plotting and
it is not very configurable. To use different plotting symbols, you
should first create and empty ordination plot with
plot(..., type="n"), and then add
text to the created
empty frame (here
... means other arguments you want to give to your
plot command). The
text commands are fully
configurable, and allow different plotting symbols and characters.
If there is a really high number of species or sites, the graphs often are congested and many labels are overwritten. It may be impossible to have complete readable graphics with some data sets. Below we give a brief overview of tricks you can use. Gavin Simpson’s blog From the bottom of the heap has a series of articles on “decluttering ordination plots” with more detailed discussion and examples.
plot(..., type="n"), if you are not satisfied with the default graph. (Here and below
...means other arguments you want to give to your
identifycommand if you do not need to see all labels.
ordilabelwhich uses non-transparent background to the text. The labels still shadow each other, but the uppermost labels are readable. Argument
prioritywill help in displaying the most interesting labels (see Decluttering blog, part 1).
orditorpfunction that uses labels only if these can be added to a graph without overwriting other labels, and points otherwise, if you do not need to see all labels. You must first create an empty plot using
plot(..., type="n"), and then add labels or points with
orditorp(see Decluttering blog).
ordipointlabelwhich uses points and text labels to the points, and tries to optimize the location of the text to minimize the overlap (see Decluttering blog).
pointsfunctions have argument
selectthat can be used for full control of selecting items plotted as text or points.
orditkplotfunction that lets you drag labels of points to better positions if you need to see all labels. Only one set of points can be used (see Decluttering blog).
plotfunctions allow you to zoom to a part of the graph using
ylimarguments to reduce clutter in congested areas.
ylim with flipped limits. If you have model
mod <- cca(dune) you can flip the first axis with
plot(mod, xlim = c(3, -2)).
You can use
ylim arguments in
ordiplot to zoom
into ordination diagrams. Normally you must set both
because ordination plots will keep the equal aspect ratio of axes, and
they will fill the graph so that the longer axis will fit.
Dynamic zooming can be done with function
orditkplot. You can directly
save the edited
orditkplot graph in various graphic formats, or you
can export the graph object back to R and use
plot to display the
No. It may be possible to port TWINSPAN to vegan, but it is not among the vegan top priorities. If anybody wants to try porting, I will be happy to help. TWINSPAN has a very permissive license, and it would be completely legal to port the function into R.
The permutation scheme influences the permutation distribution of the statistics and probably the significance levels, but does not influence the calculation of the statistics.
Some vegan functions, such as
radfit use base R facility of
family in maximum likelihood estimation. This allows use of several
alternative error distributions, among them
"gaussian". The R
family also defines the deviance. You can see the
equations for deviance with commands like
In general, deviance is 2 times log.likelihood shifted so that models with exact fit have zero deviance.