Rarefaction Species Richness
Rarefied species richness for community ecologists.
rarefy(x, sample, se = FALSE, MARGIN = 1) rrarefy(x, sample) drarefy(x, sample) rarecurve(x, step = 1, sample, xlab = "Sample Size", ylab = "Species", label = TRUE, col, lty, ...) rareslope(x, sample)
x |
Community data, a matrix-like object or a vector. |
MARGIN |
Margin for which the index is computed. |
sample |
Subsample size for rarefying community, either a single value or a vector. |
se |
Estimate standard errors. |
step |
Step size for sample sizes in rarefaction curves. |
xlab, ylab |
Axis labels in plots of rarefaction curves. |
label |
Label rarefaction curves by rownames of |
col, lty |
plotting colour and line type, see
|
... |
Parameters passed to |
Function rarefy
gives the expected species richness in random
subsamples of size sample
from the community. The size of
sample
should be smaller than total community size, but the
function will work for larger sample
as well (with a warning)
and return non-rarefied species richness (and standard error =
0). If sample
is a vector, rarefaction of all observations is
performed for each sample size separately. Rarefaction can be
performed only with genuine counts of individuals. The function
rarefy
is based on Hurlbert's (1971) formulation, and the
standard errors on Heck et al. (1975).
Function rrarefy
generates one randomly rarefied community
data frame or vector of given sample
size. The sample
can be a vector giving the sample sizes for each row. If the
sample
size is equal to or larger than the observed number
of individuals, the non-rarefied community will be returned. The
random rarefaction is made without replacement so that the variance
of rarefied communities is rather related to rarefaction proportion
than to the size of the sample
. Random rarefaction is
sometimes used to remove the effects of different sample
sizes. This is usually a bad idea: random rarefaction discards valid
data, introduces random error and reduces the quality of the data
(McMurdie & Holmes 2014). It is better to use normalizing
transformations (decostand
in vegan) possible
with variance stabilization (decostand
and
dispweight
in vegan) and methods that are not
sensitive to sample sizes.
Function drarefy
returns probabilities that species occur in
a rarefied community of size sample
. The sample
can be
a vector giving the sample sizes for each row. If the sample
is equal to or smaller than the observed number of individuals, all
observed species will have sampling probability 1.
Function rarecurve
draws a rarefaction curve for each row of
the input data. The rarefaction curves are evaluated using the
interval of step
sample sizes, always including 1 and total
sample size. If sample
is specified, a vertical line is
drawn at sample
with horizontal lines for the rarefied
species richnesses.
Function rareslope
calculates the slope of rarecurve
(derivative of rarefy
) at given sample
size; the
sample
need not be an integer.
A vector of rarefied species richness values. With a single
sample
and se = TRUE
, function rarefy
returns a
2-row matrix with rarefied richness (S
) and its standard error
(se
). If sample
is a vector in rarefy
, the
function returns a matrix with a column for each sample
size,
and if se = TRUE
, rarefied richness and its standard error are
on consecutive lines.
Function rarecurve
returns invisible
list of
rarefy
results corresponding each drawn curve.
Jari Oksanen
Heck, K.L., van Belle, G. & Simberloff, D. (1975). Explicit calculation of the rarefaction diversity measurement and the determination of sufficient sample size. Ecology 56, 1459–1461.
Hurlbert, S.H. (1971). The nonconcept of species diversity: a critique and alternative parameters. Ecology 52, 577–586.
McMurdie, P.J. & Holmes, S. (2014). Waste not, want not: Why rarefying microbiome data is inadmissible. PLoS Comput Biol 10(4): e1003531. doi: 10.1371/journal.pcbi.1003531
data(BCI) S <- specnumber(BCI) # observed number of species (raremax <- min(rowSums(BCI))) Srare <- rarefy(BCI, raremax) plot(S, Srare, xlab = "Observed No. of Species", ylab = "Rarefied No. of Species") abline(0, 1) rarecurve(BCI, step = 20, sample = raremax, col = "blue", cex = 0.6)
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