This is a maintenance release to avoid warning messages caused by changes in CRAN repository. The namespace usage is also more stringent to avoid warnings and notes in development versions of R.
vegan can be installed and loaded without tcltk
package. The tcltk package is needed in
function for interactive editing of ordination graphics.
ordisurf failed if gam package was loaded due
to namespace issues: some support functions of gam were used
instead of mgcv functions.
tolerance function failed for unconstrained
estimateR uses a more exact variance formula for
bias-corrected Chao estimate of extrapolated number of
species. The new formula may be unpublished, but it was derived
following the guidelines of Chiu, Wang, Walther & Chao,
Biometrics 70, 671–682 (2014),
Diversity accumulation functions
estaccumR use now permute package for permutations
of the order of sampling sites. Normally these functions only
need simple random permutation of sites, but restricted
permutation of the permute package and user-supplied
permutation matrices can be used.
estaccumR function can use parallel processing.
linestack accepts now expressions as labels. This
allows using mathematical symbols and formula given as
Several vegan functions can now use parallel
processing for slow and repeating calculations. All these
functions have argument
parallel. The argument can be an
integer giving the number of parallel processes. In unix-alikes
(Mac OS, Linux) this will launch
and in Windows it will set up
"snow" clusters as desribed
in the documentation of the parallel package. If
"mc.cores" is set to an integer > 1, this will be used to
automatically start parallel processing. Finally, the argument
can also be a previously set up
"snow" cluster which will
be used both in Windows and in unix-alikes. Vegan vignette
on Design decision explains the implementation (use
vegandocs("decission"), and parallel package has more
extensive documentation on parallel processing in R.
The following function use parallel processing in analysing
simper. In addition,
bioenv can compare several candidate sets of models in
metaMDS can launch several random starts in
oecosimu can evaluate test statistics for
several null models in parallel.
All permutation tests are based on the permute package
which offers strong tools for restricted permutation. All these
functions have argument
permutations. The default usage of
simple non-restricted permutations is achieved by giving a single
integer number. Restricted permutations can be defined using the
how function of the permute package. Finally, the
argument can be a permutation matrix where rows define
permutations. It is possible to use external or user constructed
help(permutations) for a brief introduction on
permutations in vegan, and permute package for the
full documention. The vignette of the permute package can
be read from vegan with command
The following functions use the permute package:
envfit (plus associated
Community null model generation has been completely
redesigned and rewritten. The communities are constructed with
nullmodel function and defined in a low level
commsim function. The actual null models are generated
simulate function that builds an array of null
models. The new null models include a wide array of quantitative
models in addition to the old binary models, and users can plug
in their own generating functions. The basic tool invoking and
analysing null models is
oecosimu. The null models are
often used only for the analysis of nestedness, but the
oecosimu allows analysing any
statistic, and null models are better seen as an alternative to
vegan package dependencies and namespace imports were adapted to changes in R, and no more trigger warnings and notes in package tests.
Three-dimensional ordination graphics using scatterplot3d for static plots and rgl for dynamic plots were removed from vegan and moved to a companion package vegan3d. The package is available in CRAN.
dispweight implements dispersion weighting
of Clarke et al. (Marine Ecology Progress Series, 320,
11–27). In addition, we implemented a new method for
generalized dispersion weighting
methods downweight species that are significantly
hclust support functions
rev are similar as these functions for
objects in base R. However,
reorder can use (and defaults
to) weighted mean. In weighted mean the node average is always the
mean of member leaves, whereas the
dendrogram uses always
unweighted means of joined branches.
ordiellipse and provides a randomization test for the
one-sided alternative hypothesis that convex hulls or ellipses in
two-dimensional ordination space have smaller areas than with
permustats extracts and inspects permutation
results with support functions
qqnorm are standard R tools that only
work with one statistic, and
are lattice graphics that work with univariate and
multivariate statistics. The results of following functions can be
permustest.cca (but not the corresponding
stressplot functions display the ordination distances
at given number of dimensions against original distances. The
method functins are similar to
metaMDS, and always use the inherent distances of each
ordination method. The functions are available for the results
cascadeKM of only one group will be
of a random value.
ordiellipse can handle points exactly on a line,
including only two points (with a warning).
radfit results for several species failed if
any of the communities had no species or had only one species.
capscale with negative
eigenvalues will now report
NA instead of using biased
simper failed when a group had only a single member.
anova.cca functions were re-written to use the
permute package. Old results may not be exactly
reproduced, and models with missing data may fail in several
cases. There is a new option of analysing a sequence of models
against each other.
simulate functions for
can return several simulations in a
object. The functions can produce simulations with correlated
errors (also for
capscale) in parametric simulation with
bioenv can use Manhattan, Gower and Mahalanobis
distances in addition to the default Euclidean. New helper
bioenvdist can extract the dissimilarities
applied in best model or any other model.
metaMDS(..., trace = 2) will show convergence
information with the default
MDSrotate can rotate a k-dimensional
ordination to k-1 variables. When these variables are
correlated (like usually is the case), the vectors can also be
correlated to previously rotated dimensions, but will be
uncorrelated to all later ones.
vegan 2.0-10 changed the weighted
so that weighted analysis of binary data was equivalent to
binary analysis. However, this broke the equivalence to the
original method. Now the function has an argument
to select the method of analysis. The problem was reported and a
fix submitted by Vanderlei Debastiani (Universidade Federal do
Rio Grande do Sul, Brasil).
can handle missing values in
ordispider can now use spatial medians instead of
rankindex can use Manhattan, Gower and Mahalanobis
distance in addition to the default Euclidean.
User can set colours and line types in function
rarecurve for plotting rarefaction curves.
spantree gained a support function
to change the minimum spanning tree into an
fitspecaccum can do weighted analysis. Gained
Functions for extrapolated number of species or for the size of species pool using Chao method were modified following Chiu et al., Biometrics 70, 671–682 (2014).
specpool can now use (and defaults to)
small sample correction with number of sites as the sample
size. Function uses basic Chao extrapolation based on the ratio of
singletons and doubletons, but switches now to bias corrected Chao
extrapolation if there are no doubletons (species found
twice). The variance formula for bias corrected Chao was derived
following the supporting
and differs slightly from Chiu et al. (2014).
poolaccum function was changed similarly, but the small
sample correction is used always.
The abundance based
estimateR uses bias corrected Chao
extrapolation, but earlier it estimated its variance with classic
Chao model. Now we use the widespread
equation for variance.
With these changes these functions are more similar to EstimateS.
tabasco uses now
hclust object for better ordering than previously when it
cast trees to
treedist default now to
match.force = TRUE and can be silenced with
verbose = FALSE.
vegdist gained Mahalanobis distance.
Nomenclature updated in plant community data with the help
of Taxonstand and taxize packages. The taxonomy of
dune data was adapted to the same sources and APG
dune use 8-character names (4
from genus + 4 from species epithet). New data set on
phylogenetic distances for
dune was extracted from Zanne
et al. (Nature 506, 89–92; 2014).
User configurable plots for
strata are deprecated in permutations. It is still
accepted but will be phased out in next releases. Use
of permute package.
capscale do not return
scores scaled by eigenvalues: use
scores function to
extract scaled results.
commsimulator is deprecated. Replace
commsimulator(x, method) with
densityplot for permutation
results are deprecated: use
permustats with its
This version is adapted to the changes in permute
package version 0.8-0 and no more triggers NOTEs in package
checks. This release may be the last of the 2.0 series, and the
next vegan release is scheduled to be a major release with
oecosimu and community pattern simulation,
support for parallel processing, and full support of the
permute package. If you are interested in these
developments, you may try the development versions of
GitHub and report the
problems and user experience to us.
envfit function assumed that all external variables
were either numeric or factors, and failed if they were, say,
character strings. Now only numeric variables are taken as
continuous vectors, and all other variables (character strings,
logical) are coerced to factors if possible. The function also
should work with degenerate data, like only one level of a
factor or a constant value of a continuous environmental
variable. The ties were wrongly in assessing permutation
nestednodf with quantitative data was not
consistent with binary models, and the fill was wrongly
calculated with quantitative data.
oecosimu now correctly adapts displayed quantiles
of simulated values to the
alternative test direction.
renyiaccum plotting failed if only one level of
scale was used.
The Kempton and Taylor algorithm was found unreliable in
fisher.alpha, and now the estimation
of Fisher alpha is only based on the number of
species and the number of individuals. The estimation of
standard errors and profile confidence intervals also had to be
tsallisaccum functions gained
renyiaccum can now add a
collector curve to
to the analysis. The collector curve is the diversity
accumulation in the order of the sampling units. With an
interesting ordering or sampling units this allows comparing
actual species accumulations with the expected randomized
specaccum can now perform weighted accumulation
using the sampling effort as weights.
This version is released due to changes in programming interface and testing procedures in R 3.0.2. If you are using an older version of R, there is no need to upgrade vegan. There are no new features nor bug fixes. The only user-visible changes are in documentation and in output messages and formatting. Because of R changes, this version is dependent on R version 2.14.0 or newer and on lattice package.
This is a maintenance release that fixes some issues raised by changed in R toolset for processing vignettes. In the same we also fix some typographic issues in the vignettes.
ordisurf gained new arguments for more flexible
definition of fitted models to better utilize the
The linewidth of contours can
now be set with the argument
Labels to arrows are positioned in a better way in
plot functions for the results of
capscale. The labels should no
longer overlap the arrow tips.
The setting test direction is clearer in
ordipointlabel gained a
plot method that can
be used to replot the saved result.
tabasco() is a new function for graphical display
of community data matrix. Technically it is an interface to R
heatmap, but its use is closer to vegan function
vegemite. The function can reorder the community data
matrix similarly as
vegemite, for instance, by ordination
heatmap, it only displays dendrograms if
supplied by the user, and it defaults to re-order the
dendrograms by correspondence analysis. Species are ordered to
match site ordering or like determined by the user.
fitspecaccum(..., model = "asymp") fitted
logistic model instead of asymptotic model (or the same as
model = "logis").
nestedtemp() failed with very sparse data (fill
plot function for constrained ordination
TRUE) which can be used
to suppress axis scale for biplot arrays.
Number of iterations in nonmetric multidimensional scaling
(NMDS) can be set with keyword
The result objects of
capscale will no longer have scores
wa.eig in the future versions of
vegan. This change does not influence normal usage,
because vegan functions do not need these items. However,
external scripts and packages may need changes in the future
versions of vegan.
The species scores were scaled wrongly in
capscale(). They were scaled correctly only when Euclidean
distances were used, but usually
capscale() is used with
non-Euclidean distances. Most graphics will change and should be
redone. The change of scaling mainly influences the spread of
species scores with respect to the site scores.
clamtest() failed to set the minimum
abundance threshold in some cases. In addition, the output was
wrong when some of the possible species groups were missing. Both
problems were reported by Richard Telford (Bergen, Norway).
Plotting an object fitted by
envfit() would fail if
p.max was used and there were unused levels for one or
more factors. The unused levels could result from deletion of
observations with missing values or simply as the result of
supplying a subset of a larger data set to
multipart() printed wrong information about the
analysis type (but did the analysis correctly). Reported by
oecosimu() failed if its
nestedfun returned a
data frame. A more fundamental fix will be in vegan 2.2-0,
where the structure of the
oecosimu() result will change.
The plot of two-dimensional
often draw original axes in a wrong angle. The problem was
reported by Elizabeth Ottesen (MIT).
treedive() for functional or phylogenetic
diversity did not correctly match the species names between the
community data and species tree when the tree contained species
that did not occur in the data. Related function
treedist() for phylogenetic distances did not try to match
the names at all.
The output of
capscale() displays the value of the
additive constant when argument
add = TRUE was used.
fitted() functions for
capscale() can now return conditioned (partial) component
of the response: Argument
model gained a new alternative
model = "pCCA".
dispindmorisita() output gained a new column for
Chi-squared based probabilities that the null hypothesis (random
distribution) is true.
monoMDS() have new default
convergence criteria. Most importantly, scale factor of the
sfgrmin) is stricter. The former limit was too
slack with large data sets and iterations stopped early without
getting close to the solution. In addition,
ignore now requests to dimensions beyond those calculated
instead of failing, and
results do not drop dimensions.
legend argument for
positioning the legend.
nestednodf() gained a
ordiR2step() gained new argument
TRUE) which can be used to turn off the criterion
of stopping when the adjusted R-squared of the current
model exceeds that of the scope. This option allows model
building when the
scope would be overdetermined (number of
predictors higher than number of observations).
ordiR2step() now handles partial redundancy analysis
orditorp() gained argument
select to select
the rows or columns of the results to display.
protest() prints the standardized residual statistic
squared m12 in addition to the squared Procrustes
correlation R-squared. Both were calculated, but only
the latter was displayed.
Permutation tests are much faster in
of calling repeatedly
procrustes(), the goodness of fit
statistic is evaluated within the function.
wcmdscale() gained methods for
plot etc. of the results. These methods are only used if
wcmdscale result is returned with, e.g., argument
eig = TRUE. The default is still to return only a matrix of
scores similarly as the standard R function
and in that case the new methods are not used.
anova(<cca_object>, ...) failed with
by = "axis" and
by = "term". The bug was reported by
Dr Sven Neulinger (Christian Albrecht University, Kiel, Germany).
radlattice did not honour argument
BIC = TRUE,
but always displayed AIC.
Most vegan functions with permutation tests have now a
density method that can be used to find empirical
probability distributions of permutations. There is a new
plot method for these functions that displays both the
density and the observed statistic. The
density function is
adonis can return several statistics, and it has
densityplot method (based on lattice).
oecosimu already had
densityplot, but they are now similar to other vegan
methods, and also work with
radfit functions got a
predict method that
also accepts arguments
total for new
ranks and site totals for prediction. The functions can also
interpolate to non-integer “ranks”, and in some models
Labels can now be set in the
results. The labels must be given in the same order that the
function uses internally, and new support function
can be used to display the default labels in their correct order.
Mantel tests (functions
mantel.partial) gained argument
na.rm which can be
used to remove missing values. This options should be used with
care: Permutation tests can be biased if the missing values were
originally in matching or fixed positions.
radfit results can be consistently accessed with
the same methods whether they were a single model for a single
site, all models for a single site or all models for all sites
in the data. All functions now have methods
Building of vegan vignettes failed with the latest version of LaTeX (TeXLive 2012).
R versions later than 2.15-1 (including development
version) report warnings and errors when installing and checking
vegan, and you must upgrade vegan to this version.
The warnings concern functions
betadisper, and the error occurs in
These errors and warnings were triggered by internal changes in
adipart assumed constant gamma diversity in
simulations when assessing the P-value. This could give
biased results if the null model produces variable gamma
diversities and option
weights = "prop" is used. The
default null model (
"r2dtable") and the default option
weights = "unif") were analysed correctly.
anova(<prc-object>, by = "axis") and other
by cases failed due to ‘NAMESPACE’ issues.
clamtest wrongly used frequencies instead of the
counts when calculating sample coverage. No detectable
differences were produced when rerunning examples from Chazdon
et al. 2011 and vegan help page.
envfit failed with unused factor levels.
cca results with
type = "response" or
type = "working" failed with
newdata if the number of rows did not match with the
original data. Now the
newdata is ignored if it has a
wrong number of rows. The number of rows must match because
the results in
cca must be weighted by original row
totals. The problem did not concern
capscale results which do not need row weights.
Reported by Glenn De'ath.
Functions for diversity partitioning (
multipart) have now
default methods. The
formula method is
identical to the previous functions, but the
method can take two matrices as input.
multipart can be used for
fast and easy overall partitioning to alpha, beta and gamma
diversities by omitting the argument describing the hierarchy.
The method in
betadisper is biased with small
sample sizes. The effects of the bias are strongest with
unequal sample sizes. A bias adjusted version was developed by
Adrian Stier and Ben Bolker, and can be invoked with argument
bias.adjust (defaults to
bioenv accepts dissimilarities (or square matrices
that can be interpreted as dissimilarities) as an alternative to
community data. This allows using other dissimilarities than
those available in
plot function for
envfit results gained new
bg that can be used to set background colour for
msoplot is more configurable, and allows, for
instance, setting y-axis limits.
Hulls and ellipses are now filled using semitransparent
ordiellipse, and the
user can set the degree of transparency with a new argument
alpha. The filled shapes are used when these functions
are called with argument
draw = "polygon". Function
ordihull puts labels (with argument
label = TRUE)
now in the real polygon centre.
ordiplot3d returns function
and the projected location of the
these can be used to add
envfit results to existing
Equal aspect ratio cannot be set exactly in
because underlying core routines do not allow this. Now
ordiplot3d sets equal axis ranges, and the documents
urge users to verify that the aspect ratio is reasonably equal
and the graph looks like a cube. If the problems cannot be
solved in the future,
ordiplot3d may be removed from
next releases of vegan.
ordipointlabel gained argument to
select only some of the items for plotting. The
argument can be used only with one set of points.
Added new nestedness functions
nestedbetajac that implement multiple-site dissimilarity
indices and their decomposition into turnover and nestedness
components following Baselga (Global Ecology and
Biogeography 19, 134–143; 2010).
rarecurve to draw rarefaction curves
for each row (sampling unit) of the input data, optionally with
lines showing rarefied species richness with given sample size
for each curve.
simper that implements
“similarity percentages” of Clarke (Australian
Journal of Ecology 18, 117–143; 1993). The method compares
two or more groups and decomposes the average between-group
Bray-Curtis dissimilarity index to contributions by individual
species. The code was developed in
by Eduard Szöcs (Uni Landau, Germany).
betadisper() failed when the
groups was a
factor with empty levels.
Some constrained ordination methods and their support
functions are more robust in border cases (completely aliased
effects, saturated models, user requests for non-existng scores
plot function for constrained ordination, and
anova(<cca.object>, by = "margin").
scores function for
monoMDS did not
choices argument and hence dimensions could not be
scores method failed if the number of
requested axes was higher than the ordination object had. This
was reported as an error in
R-sig-ecology mailing list.
noshare = 0 is now
regarded as a numeric threshold that always triggers extended
stepacross), instead of being treated
as synonymous with
noshare = FALSE which always
suppresses extended dissimilarities.
Nestedness discrepancy index
nesteddisc gained a
new argument that allows user to set the number of iterations
in optimizing the index.
oecosimu displays the mean of simulations and
describes alternative hypothesis more clearly in the printed
Implemented adjusted R-squared for partial
RDA. For partial model
rda(Y ~ X1 + Condition(X2)) this
is the same as the component
[a] = X1|X2 in variance
varpart and describes the marginal (unique)
effect of constraining term to adjusted R-squared.
Added Cao dissimilarity (CYd) as a new dissimilarity
vegdist following Cao et al., Water
Envir Res 69, 95–106 (1997). The index should be good for
data with high beta diversity and variable sampling
intensity. Thanks to consultation to Yong Cao (Univ Illinois,
capscale failed if constrained component
had zero rank. This happened most likely in partial models when
the conditions aliased constraints. The problem was observed in
anova(..., by ="margin") which uses partial models to
analyses the marginal effects, and was reported in an email
goodness sometimes failed when
metaMDS was based on
isoMDS (MASS package)
metaMDSdist did not use the same defaults for
step-across (extended) dissimilarities as
metaMDS(..., engine = "isoMDS"). The change of defaults can
also influence triggering of step-across in
capscale(..., metaMDSdist = TRUE).
adonis contained a minor bug resulting from
incomplete implementation of a speed-up that did not affect the
results. In fixing this bug, a further bug was identified in
transposing the hat matrices. This second bug was only active
following fixing of the first bug. In fixing both bugs, a
speed-up in the internal f.test() function is fully
realised. Reported by Nicholas Lewin-Koh.
ordisegments gained argument
order.by that gives a variable to sort points within
groups. Earlier the points were assumed to be in order.
ordispider invisibly returns the
coordinates to which the points were connected. Typically these
are class centroids of each point, but for constrained ordination
groups they are the LC scores.
clamtest: new function to classify species as
generalists and specialists in two distinct habitats (CLAM test of
Chazdon et al., Ecology 92, 1332–1343; 2011). The test is
based on multinomial distribution of individuals in two habitat
types or sampling units, and it is applicable only to count data
with no over-dispersion.
plot method (which
also can add items to existing plots). These are similar as
fisherfit, but display only data without the fitted lines.
raupcrick: new function to implement Raup-Crick
dissimilarity as a probability of number of co-occurring species
with occurrence probabilities proportional to species
frequencies. Vegan has Raup-Crick index as a choice in
vegdist, but that uses equal sampling probabilities for
species and analytic equations. The new
function uses simulation with
oecosimu. The function
follows Chase et al. (2011) Ecosphere 2:art24
and was developed with the consultation of Brian Inouye.
meandist could scramble items and give
wrong results, especially when the
numerical. The problem was reported by Dr Miguel Alvarez
metaMDS did not reset
tries when a new model
was started with a
previous.best solution from a different
permatswap for community null models using
quantitative swap never swapped items in a 2 by 2
submatrix if all cells were filled.
The result from
permutest.cca could not be
updated because of a ‘NAMESPACE’ issue.
R 2.14.0 changed so that it does not accept using
sd() function for matrices (which was the behaviour at
least since R 1.0-0), and several vegan functions were
changed to adapt to this change (
simulate methods for
capscale). The change in R 2.14.0 does not influence the
results but you probably wish to upgrade vegan to avoid
nesteddisc is slacker and hence faster when trying
to optimize the statistic for tied column frequencies. Tracing
showed that in most cases an improved ordering was found rather
early in tries, and the results are equally good in most cases.
Peter Minchin joins the vegan team.
vegan implements standard R ‘NAMESPACE’. In
S3 methods are not exported which means that you
cannot directly use or see contents of functions like
use these functions you should rely on R delegation and simply
cca and for its result objects use
anova without suffix
.cca. To see the contents of
the function you can use
:::, such as
vegan:::cca.default. This change may break packages,
documents or scripts that rely on non-exported names.
vegan depends on the permute package. This
package provides powerful tools for restricted permutation
schemes. All vegan permutation will gradually move to use
permute, but currently only
betadisper uses the new
monoMDS: a new function for non-metric
multidimensional scaling (NMDS). This function replaces
MASS::isoMDS as the default method in
metaMDS. Major advantages of
monoMDS are that it
has ‘weak’ (‘primary’) tie treatment which means
that it can split tied observed dissimilarities. ‘Weak’
tie treatment improves ordination of heterogeneous data sets,
because maximum dissimilarities of 1 can be split. In
addition to global NMDS,
monoMDS can perform local and
hybrid NMDS and metric MDS. It can also handle missing and zero
monoMDS is faster than
previous alternatives. The function uses
written by Peter Minchin.
MDSrotate a new function to replace
metaMDSrotate. This function can rotate both
monoMDS results so that the first axis is parallel to
an environmental vector.
eventstar finds the minimum of the evenness profile
on the Tsallis entropy, and uses this to find the corresponding
values of diversity, evenness and numbers equivalent following
Mendes et al. (Ecography 31, 450-456; 2008). The code was
contributed by Eduardo Ribeira Cunha and Heloisa Beatriz Antoniazi
Evangelista and adapted to vegan by Peter Solymos.
fitspecaccum fits non-linear regression models to
the species accumulation results from
function can use new self-starting species accumulation models
in vegan or other self-starting non-linear regression
models in R. The function can fit Arrhenius, Gleason, Gitay,
Lomolino (in vegan), asymptotic, Gompertz,
Michaelis-Menten, logistic and Weibull (in base R) models. The
Self-starting non-linear species accumulation models
SSlomolino. These can be used with
directly in non-linear regression with
nls. These functions
were implemented because they were found good for species-area
models by Dengler (J. Biogeogr. 36, 728-744; 2009).
mrpp warn on negative dissimilarities, and
betadisper refuses to analyse them. All these functions
expect dissimilarities, and giving something else (like
correlations) probably is a user error.
betadisper uses restricted permutation of the
monoMDS as its default ordination
engine. Function gains new argument
engine that can be used
to alternatively select
MASS::isoMDS. The default is not
monoMDS because its
‘weak’ tie treatment can cope with tied maximum
dissimilarities of one. However,
stepacross is the default
isoMDS because it cannot handle adequately these tied
predict method which uses
either linear or spline interpolation for data between observed
points. Extrapolation is possible with spline interpolation, but
may make little sense.
specpool can handle missing values or empty factor
levels in the grouping factor
pool. Now also checks that
the length of the
pool matches the number of
metaMDSrotate was replaced with
that can also handle the results of
permuted.index2 and other “new” permutation
code was removed in favour of the permute package. This code
was not intended for normal use, but packages depending on that
code in vegan should instead depend on permute.
treeheight uses much snappier code. The results
should be unchanged.