| Title: | Integrating Phylogenies and Ecology |
|---|---|
| Description: | Functions for phylocom integration, community analyses, null-models, traits and evolution. Implements numerous ecophylogenetic approaches including measures of community phylogenetic and trait diversity, phylogenetic signal, estimation of trait values for unobserved taxa, null models for community and phylogeny randomizations, and utility functions for data input/output and phylogeny plotting. A full description of package functionality and methods are provided by Kembel et al. (2010) <doi:10.1093/bioinformatics/btq166>. |
| Authors: | Steven W. Kembel <[email protected]>, David D. Ackerly <[email protected]>, Simon P. Blomberg <[email protected]>, Will K. Cornwell <[email protected]>, Peter D. Cowan <[email protected]>, Matthew R. Helmus <[email protected]>, Helene Morlon <[email protected]>, Campbell O. Webb <[email protected]> |
| Maintainer: | Steven W. Kembel <[email protected]> |
| License: | GPL-2 |
| Version: | 1.8.3 |
| Built: | 2024-11-10 03:44:14 UTC |
| Source: | https://github.com/skembel/picante |
Functions for phylocom integration, community analyses, null-models, traits and evolution. Implements numerous ecophylogenetic approaches including measures of community phylogenetic and trait diversity, phylogenetic signal, estimation of trait values for unobserved taxa, null models for community and phylogeny randomizations, and utility functions for data input/output and phylogeny plotting. A full description of package functionality and methods are provided by Kembel et al. (2010) <doi:10.1093/bioinformatics/btq166>.
| Package: | picante |
| Type: | Package |
| Version: | 1.8.3 |
| Date: | 2023-07-10 |
| License: | GPL-2 |
Author: Steven W. Kembel <[email protected]>, David D. Ackerly <[email protected]>, Simon P. Blomberg <[email protected]>, Will K. Cornwell <[email protected]>, Peter D. Cowan <[email protected]>, Matthew R. Helmus <[email protected]>, Helene Morlon <[email protected]>, Campbell O. Webb <[email protected]> Maintainer: Steven W. Kembel <[email protected]>
Plots a phylogeny with tip labels colored to indicate continuous or discrete trait values
color.plot.phylo(phylo, df, trait, taxa.names, num.breaks = ifelse(is.factor(df[,trait]), length(levels(df[,trait])), 12), col.names = rainbow(ifelse(length(num.breaks) > 1, length(num.breaks) - 1, num.breaks)), cut.labs = NULL, leg.title = NULL, main = trait, leg.cex = 1, tip.labs = NULL, ...)color.plot.phylo(phylo, df, trait, taxa.names, num.breaks = ifelse(is.factor(df[,trait]), length(levels(df[,trait])), 12), col.names = rainbow(ifelse(length(num.breaks) > 1, length(num.breaks) - 1, num.breaks)), cut.labs = NULL, leg.title = NULL, main = trait, leg.cex = 1, tip.labs = NULL, ...)
phylo |
An object of class |
df |
A dataframe containing the traits to be plotted |
trait |
A string representing the name of column in the dataframe to be plotted |
taxa.names |
A string representing the name of column in the dataframe that contains the names of the taxa |
num.breaks |
For continuous traits, the number of bins to separate the data into |
col.names |
A vector of colors to use for tip labels |
leg.title |
A title for the tip color legend |
main |
A main title for the plot |
cut.labs |
A main title for the plot |
leg.cex |
A main title for the plot |
tip.labs |
A main title for the plot |
... |
Additional argument to pass to the |
If if trait is a factor then each level of the factor is plotted with the corresponding col.names value (if length(num.breaks) > length(col.names) colors are recycled.) If trait is not a factor then it is assumed to be continuous and trait is evenly divided into num.breaks levels.
The command is invoked for its side effect, a plot of the phylo with tips colored based on trait
Peter Cowan <[email protected]>
Calculates MPD (mean pairwise distance) separating taxa in two communities, a measure of phylogenetic beta diversity
comdist(comm, dis, abundance.weighted = FALSE)comdist(comm, dis, abundance.weighted = FALSE)
comm |
Community data matrix |
dis |
Interspecific distance matrix |
abundance.weighted |
Should mean pairwise distances separating species in two communities be weighted by species abundances? (default = FALSE) |
This function calculates a measure of phylogenetic beta diversity: the expected phylogenetic distance separating two individuals or taxa drawn randomly from different communities.
Distance object of MPD values separating each pair of communities.
Steven Kembel <[email protected]>
C.O. Webb, D.D. Ackerly, and S.W. Kembel. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Bioinformatics 18:2098-2100.
data(phylocom) comdist(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=TRUE)data(phylocom) comdist(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=TRUE)
Calculates MNTD (mean nearest taxon distance) separating taxa in two communities, a measure of phylogenetic beta diversity
comdistnt(comm, dis, abundance.weighted = FALSE, exclude.conspecifics = FALSE)comdistnt(comm, dis, abundance.weighted = FALSE, exclude.conspecifics = FALSE)
comm |
Community data matrix |
dis |
Interspecific distance matrix |
abundance.weighted |
Should mean nearest taxon distances from each species to species in the other community be weighted by species abundance? (default = FALSE) |
exclude.conspecifics |
Should conspecific taxa in different communities be exclude from MNTD calculations? (default = FALSE) |
This metric has also been referred to as MNND (mean nearest neighbour distance).
This function calculates a measure of phylogenetic beta diversity: the average phylogenetic distance to the most similar taxon or individual in the other community for taxa or individuals in two communities.
Distance object of MNTD values separating each pair of communities.
Steven Kembel <[email protected]>
C.O. Webb, D.D. Ackerly, and S.W. Kembel. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Bioinformatics 18:2098-2100.
data(phylocom) comdistnt(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=FALSE)data(phylocom) comdistnt(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=FALSE)
Calculates measures of community phylogenetic structure (correlation between co-occurrence and phylogenetic distance) to patterns expected under various null models
comm.phylo.cor(samp, phylo, metric = c("cij", "checkerboard", "jaccard", "doij"), null.model = c("sample.taxa.labels", "pool.taxa.labels", "frequency", "richness", "independentswap","trialswap"), runs = 999, ...)comm.phylo.cor(samp, phylo, metric = c("cij", "checkerboard", "jaccard", "doij"), null.model = c("sample.taxa.labels", "pool.taxa.labels", "frequency", "richness", "independentswap","trialswap"), runs = 999, ...)
samp |
Community data matrix |
phylo |
Phylogenetic tree |
metric |
Metric of co-occurrence to use (see |
null.model |
Null model to use (see Details section for description) |
runs |
Number of runs (randomizations) |
... |
Additional arguments to randomizeMatrix |
Currently implemented null models (arguments to null.model):
Shuffle phylogeny tip labels (only within set of taxa present in community data)
Shuffle phylogeny tip labels (across all taxa included in phylogenetic tree)
Randomize community data matrix abundances within species (maintains species occurence frequency)
Randomize community data matrix abundances within samples (maintains sample species richness)
Randomize community data matrix maintaining species occurrence frequency and site richnessing using independent swap
Randomize community data matrix maintaining species occurrence frequency and site richnessing using trial swap
A list with elements:
obs.corr |
Observed co-occurrence/phylogenetic distance correlation |
obs.corr.p |
P-value of observed correlation (standard P-value for correlation coefficient, not based on comparison with randomizations) |
obs.rank |
Rank of observed correlation vs. random |
runs |
Number of runs (randomizations) |
obs.rand.p |
P-value of observed correlation vs. randomizations (= obs.rank / (runs + 1)) |
random.corrs |
A vector of random correlation calculated for each run |
Steven Kembel <[email protected]>
Cavender-Bares J., D.A. Ackerly, D. Baum and F.A. Bazzaz. 2004. Phylogenetic overdispersion in Floridian oak communities, American Naturalist, 163(6):823-843.
data(phylocom) comm.phylo.cor(phylocom$sample, phylocom$phylo, metric="cij",null.model="sample.taxa.labels")data(phylocom) comm.phylo.cor(phylocom$sample, phylocom$phylo, metric="cij",null.model="sample.taxa.labels")
Calculates measures of community phylogenetic structure (quantile regression between co-occurrence and phylogenetic distance) to patterns expected under various null models
comm.phylo.qr(samp, phylo, metric = c("cij", "checkerboard", "jaccard", "doij"), null.model = c("sample.taxa.labels", "pool.taxa.labels", "frequency", "richness", "independentswap","trialswap"), quant = 0.75, runs = 999, show.plot = FALSE, ...)comm.phylo.qr(samp, phylo, metric = c("cij", "checkerboard", "jaccard", "doij"), null.model = c("sample.taxa.labels", "pool.taxa.labels", "frequency", "richness", "independentswap","trialswap"), quant = 0.75, runs = 999, show.plot = FALSE, ...)
samp |
Community data matrix |
phylo |
Phylogenetic tree |
metric |
Metric of co-occurrence to use (see |
null.model |
Null model to use (see Details section for description) |
quant |
Quantile of slope to be fit (using |
runs |
Number of runs (randomizations) |
show.plot |
Option to display a plot of co-occurrence versus phylogenetic distance with quantile regression slope fit |
... |
Additional arguments to randomizeMatrix |
This function fits a quantile regression of co-occurrence versus phylogenetic distances separating species, and compares observed patterns to the patterns expected under some null model. The quantile regressions are fit using the rq function from the quantreg package.
Currently implemented null models (arguments to null.model):
Shuffle phylogeny tip labels (only within set of taxa present in community data)
Shuffle phylogeny tip labels (across all taxa included in phylogenetic tree)
Randomize community data matrix abundances within species (maintains species occurence frequency)
Randomize community data matrix abundances within samples (maintains sample species richness)
Randomize community data matrix maintaining species occurrence frequency and site richnessing using independent swap
Randomize community data matrix maintaining species occurrence frequency and site richnessing using trial swap
A list with elements:
obs.qr.intercept |
Observed co-occurrence/phylogenetic distance quantile regression intercept |
obs.qr.slope |
Observed co-occurrence/phylogenetic distance quantile regression slope |
obs.qr.slope.p |
P-value of observed quantile regression slope significance versus null model (calculated based on comparison with randomizations) |
obs.rank |
Rank of observed quantile regression slope vs. random |
runs |
Number of runs (randomizations) |
random.qr.slopes |
A vector of quantile regression slopes calculated for each randomization |
Steven Kembel <[email protected]>
Cavender-Bares J., D.A. Ackerly, D. Baum and F.A. Bazzaz. 2004. Phylogenetic overdispersion in Floridian oak communities, American Naturalist, 163(6):823-843. Slingsby, J. A. and G. A. Verboom. 2006. Phylogenetic relatedness limits coexistence at fine spatial scales: evidence from the schoenoid sedges (Cyperaceae: Schoeneae) of the Cape Floristic Region, South Africa. The American Naturalist 168:14-27.
data(phylocom) comm.phylo.qr(phylocom$sample, phylocom$phylo, metric="cij", null.model="sample.taxa.labels", runs=99)data(phylocom) comm.phylo.qr(phylocom$sample, phylocom$phylo, metric="cij", null.model="sample.taxa.labels", runs=99)
Table of correlations with associated P-values and df, can be used with regular or independent contrast data
cor.table(x, cor.method = c("pearson","spearman"), cor.type=c("standard","contrast"))cor.table(x, cor.method = c("pearson","spearman"), cor.type=c("standard","contrast"))
x |
Data frame of data points or contrasts at nodes |
cor.method |
Correlation method (as |
cor.type |
Are data |
r |
Correlation values |
df |
Degrees of freedom |
P |
P-values |
Steven Kembel <[email protected]>
Garland, T., Jr., P. H. Harvey, and A. R. Ives. 1992. Procedures for the analysis of comparative data using phylogenetically independent contrasts. Systematic Biology 41:18-32.
Calculates evolutionary distinctiveness measures for a suite of species by: a) equal splits (Redding and Mooers 2006) b) fair proportions (Isaac et al., 2007). Returns a datafram with species identifiers and species scores.
evol.distinct(tree, type = c("equal.splits", "fair.proportion"), scale = FALSE, use.branch.lengths = TRUE)evol.distinct(tree, type = c("equal.splits", "fair.proportion"), scale = FALSE, use.branch.lengths = TRUE)
tree |
an object of class phylo |
type |
a) equal splits (Redding and Mooers 2006) or b) fair proportions (Isaac et al., 2007) |
scale |
The scale option refers to whether or not the phylogeny should be scaled to a depth of 1 or, in the case of an ultrametric tree, scaled such that branch lengths are relative. |
use.branch.lengths |
If use.branch.lengths=FALSE, then all branch lengths are changed to 1. |
This function will return a vector of evolutionary distinctivenss for every species in the given tree. If only a subset of values are needed there are two, concetually distinct options: either prune the tree first and then pass the tree in or subset the resulting vector. These two options will provide very different outputs.
Karen Magnuson-Ford, Will Cornwell, Arne Mooers, Mark Vellend
Redding, D.W. and Mooers, A.O. (2006). Incorporating evolutionary measures into conservation prioritisation. Conservation Biology, 20, 1670-1678.
Isaac, N.J.B., Turvey, S.T., Collen, B., Waterman, C. and Baillie, J.E.M. (2007). Mammals on the EDGE: conservation priorities based on threat and phylogeny. PLoS ONE, 2, e296.
Mark Vellend, William K. Cornwell, Karen Magnuson-Ford, and Arne Mooers. In press. Measuring phylogenetic biodiversity. In: Biological diversity: frontiers in measurement and assessment. Edited by Anne Magurran and Brian McGill.
Calculates the expected phylogenetic diversity (Faith's PD) and variance of PD under binomial sampling with a fixed probability of each tip being sampled, and the Edge-length Abundance Distribution of a phylogeny.
expected.pd(phy) variance.pd(phy, upper.bound=TRUE) ead(phy)expected.pd(phy) variance.pd(phy, upper.bound=TRUE) ead(phy)
phy |
phylo object |
upper.bound |
Calculate upper bound of PD variance? (default = TRUE) |
The function expected.pd calculates the expected phylogenetic diversity (Faith's PD - total branch length) for all subsets of a phylogeny, based on an analytic solution for expected PD.
The function variance.pd additionally calculates the variance of expected PD for all subsets of a phylogeny, based on an analytic solution for expected PD. If argument upper.bound=TRUE, a fast solution for the upper bound of the variance is returned. Otherwise, the exact solution for the variance is returned. Note that the exact solution is much slower than the upper bound solution.
The function ead calculates the edge abundance distribution (EAD), the length of edges with different numbers of descendant tips.
n |
Expected Number of tips sampled |
expected.pd |
Expected PD for a given n |
variance.pd |
Variance of PD for a given n |
num.children |
Number of tips descended from an edge |
edge.length |
Total phylogenetic edge length for a given number of tips descended from an edge |
Steven Kembel <[email protected]> and James O'Dwyer <[email protected]>
J.P. O'Dwyer, S.W. Kembel, and J.L. Green. 2012. Phylogenetic Diversity Theory Sheds Light on the Structure of Microbial Communities. PLoS Comput Biol 8(12): e1002832.
randtree <- rcoal(300) randtree.pd.ub <- variance.pd(randtree, upper.bound=TRUE) randtree.pd.exact <- variance.pd(randtree, upper.bound=FALSE) plot(expected.pd(randtree), xlab="Number of tips", ylab="Phylogenetic diversity (PD)", type="l", log="xy") lines(randtree.pd.exact$expected.pd+1.96*sqrt(randtree.pd.exact$variance.pd), lty=2) lines(randtree.pd.exact$expected.pd-1.96*sqrt(randtree.pd.exact$variance.pd), lty=2) lines(randtree.pd.ub$expected.pd+1.96*sqrt(randtree.pd.ub$variance.pd), lty=3) lines(randtree.pd.ub$expected.pd-1.96*sqrt(randtree.pd.ub$variance.pd), lty=3) legend("bottomright", lty=c(1,2,3), legend=c("Expected PD", "95 percent CI (exact)","95 percent CI (upper bound)"))randtree <- rcoal(300) randtree.pd.ub <- variance.pd(randtree, upper.bound=TRUE) randtree.pd.exact <- variance.pd(randtree, upper.bound=FALSE) plot(expected.pd(randtree), xlab="Number of tips", ylab="Phylogenetic diversity (PD)", type="l", log="xy") lines(randtree.pd.exact$expected.pd+1.96*sqrt(randtree.pd.exact$variance.pd), lty=2) lines(randtree.pd.exact$expected.pd-1.96*sqrt(randtree.pd.exact$variance.pd), lty=2) lines(randtree.pd.ub$expected.pd+1.96*sqrt(randtree.pd.ub$variance.pd), lty=3) lines(randtree.pd.ub$expected.pd-1.96*sqrt(randtree.pd.ub$variance.pd), lty=3) legend("bottomright", lty=c(1,2,3), legend=c("Expected PD", "95 percent CI (exact)","95 percent CI (upper bound)"))
Data on the structure of a host-parasitoid food web from Ives & Godfray (2006). Includes information on phylogenetic covariances among 12 leaf-mining moth hosts and 27 species of parasitoid wasps.
data(IvesGodfray)data(IvesGodfray)
A list with three elements:
host Phylogenetic variance/covariance matrix for 12 leaf-mining moth hosts
parasitoid Phylogenetic variance/covariance matrix for 27 species of parasitoid wasps
interactions Matrix describing interactions between hosts and parasitoids
Ives A.R. & Godfray H.C. (2006) Phylogenetic analysis of trophic associations. The American Naturalist, 168, E1-E14
Calculates K statistic of phylogenetic signal
Kcalc(x, phy, checkdata=TRUE)Kcalc(x, phy, checkdata=TRUE)
x |
Vector or data.frame of trait data (in phylo\$tip.label order) |
phy |
phylo object |
checkdata |
Check for match between trait and phylogeny taxa labels using |
K |
K statistic |
Simon Blomberg <[email protected]> and David Ackerly <[email protected]>
Blomberg, S. P., and T. Garland, Jr. 2002. Tempo and mode in evolution: phylogenetic inertia, adaptation and comparative methods. Journal of Evolutionary Biology 15:899-910.
Blomberg, S. P., T. Garland, Jr., and A. R. Ives. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717-745.
randtree <- rcoal(20) randtraits <- rTraitCont(randtree) Kcalc(randtraits[randtree$tip.label],randtree)randtree <- rcoal(20) randtraits <- rTraitCont(randtree) Kcalc(randtraits[randtree$tip.label],randtree)
These functions compare taxa present in phylogenies with community or trait data sets, pruning and sorting the two kinds of data to match one another for subsequent analysis.
match.phylo.comm(phy, comm) match.phylo.data(phy, data) match.comm.dist(comm, dis)match.phylo.comm(phy, comm) match.phylo.data(phy, data) match.comm.dist(comm, dis)
phy |
A phylogeny object of class phylo |
comm |
Community data matrix |
data |
A data object - a vector (with names matching phy) or a data.frame or matrix (with row names matching phy) |
dis |
A distance matrix - a dist or matrix object |
A common pitfall in comparative analyses in R is that taxa labels are assumed to match between phylogenetic and other data sets. These functions prune a phylogeny and community or trait data set to match one another, reporting taxa that are missing from one data set or the other.
Taxa names for phylogeny objects are taken from the phylogeny's tip labels. Taxa names for community data are taken from the column names. Taxa names for trait data are taken from the element names (vector) or row names (data.frame or matrix). Taxa names for distance data are taken from column/row names of the distance matrix/dist object.
If community data lack taxa names, the function will issue a warning and no result will be returned, since the community-phylogenetic analyses in picante require named taxa in the community data set.
If trait data or distance matrix lack names, a warning is issued and the data are assumed to be sorted in the same order as the phylogeny's tip labels or community's column labels.
These utility functions are used by several functions that assume taxa labels in phylogeny and data match, including Kcalc, phylosignal, and raoD.
A list containing the following elements, pruned and sorted to match one another:
phy |
A phylogeny object of class phylo |
comm |
Community data matrix |
data |
A data object (vector, data.frame or matrix) |
dist |
A distance matrix - a dist or matrix object |
Steven Kembel <[email protected]>
data(phylocom) match.phylo.comm(phylocom$phylo, phylocom$sample) match.phylo.data(phylocom$phylo, phylocom$traits[1:10,])data(phylocom) match.phylo.comm(phylocom$phylo, phylocom$sample) match.phylo.data(phylocom$phylo, phylocom$traits[1:10,])
Converts a community data matrix to a Phylocom database-format community sample
matrix2sample(z)matrix2sample(z)
z |
Community data matrix |
Phylocom database-format community sample
Steven Kembel <[email protected]> and Cam Webb <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
data(phylocom) matrix2sample(phylocom$sample)data(phylocom) matrix2sample(phylocom$sample)
Calculates MNTD (mean nearest taxon distance) for taxa in a community
mntd(samp, dis, abundance.weighted=FALSE)mntd(samp, dis, abundance.weighted=FALSE)
samp |
Community data matrix |
dis |
Interspecific distance matrix |
abundance.weighted |
Should mean nearest taxon distances for each species be weighted by species abundance? (default = FALSE) |
This metric has also been referred to as MNND (mean nearest neighbour distance), and the function was named mnnd in picante versions < 0.7.
Vector of MNTD values for each community.
Steven Kembel <[email protected]>
Webb, C., D. Ackerly, M. McPeek, and M. Donoghue. 2002. Phylogenies and community ecology. Annual Review of Ecology and Systematics 33:475-505.
data(phylocom) mntd(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=TRUE)data(phylocom) mntd(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=TRUE)
Calculates mean pairwise distance separating taxa in a community
mpd(samp, dis, abundance.weighted=FALSE)mpd(samp, dis, abundance.weighted=FALSE)
samp |
Community data matrix |
dis |
Interspecific distance matrix |
abundance.weighted |
Should mean pairwise distances be weighted by species abundance? (default = FALSE) |
Vector of MPD values for each community
Steven Kembel <[email protected]>
Webb, C., D. Ackerly, M. McPeek, and M. Donoghue. 2002. Phylogenies and community ecology. Annual Review of Ecology and Systematics 33:475-505.
data(phylocom) mpd(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=TRUE)data(phylocom) mpd(phylocom$sample, cophenetic(phylocom$phylo), abundance.weighted=TRUE)
Calculates phylogenetic signal for data.frame of traits. Traits may have missing values in which case the tree will be pruned prior to calculating phylogenetic signal for each trait.
multiPhylosignal(x, phy, checkdata=TRUE, ...)multiPhylosignal(x, phy, checkdata=TRUE, ...)
x |
Data frame of trait data (traits in columns) with row names corresponding to tip.labels |
phy |
phylo object |
checkdata |
Check for match between trait and phylogeny taxa labels using |
... |
Additional arguments to phylosignal |
Returns a data frame with phylogenetic signal results for each trait
Steven Kembel <[email protected]>
Fits a linear model to the association strengths of a bipartite data set with or without phylogenetic correlation among the interacting species
pblm(assocs, tree1=NULL, tree2=NULL, covars1=NULL, covars2=NULL, bootstrap=FALSE, nreps=10, maxit=10000, pstart=c(.5,.5)) pblmpredict(x, tree1.w.novel=NULL, tree2.w.novel=NULL, predict.originals=FALSE)pblm(assocs, tree1=NULL, tree2=NULL, covars1=NULL, covars2=NULL, bootstrap=FALSE, nreps=10, maxit=10000, pstart=c(.5,.5)) pblmpredict(x, tree1.w.novel=NULL, tree2.w.novel=NULL, predict.originals=FALSE)
assocs |
A matrix of association strengths among two sets of interacting species |
tree1 |
A phylo tree object or a phylogenetic covariance matrix for the rows of |
tree2 |
A phylo tree object or a phylogenetic covariance matrix for the columns of |
covars1 |
A matrix of covariates (e.g., traits) for the row species of |
covars2 |
A matrix of covariates (e.g., traits) for the column species of |
bootstrap |
logical, bootstrap confidence intervals of the parameter estimates |
nreps |
Number of bootstrap replicated data sets to estimate parameter CIs |
maxit |
as in |
pstart |
starting values of the two phylogenetic signal strength parameters passed to |
x |
object of class |
tree1.w.novel |
A phylo tree object or a phylogenetic covariance matrix which corresponds to |
tree2.w.novel |
A phylo tree object or a phylogenetic covariance matrix which corresponds to |
predict.originals |
if |
Fit a linear model with covariates using estimated generalized least squares to the association strengths between two sets of interacting species.
Associations can be either binary or continuous. If phylogenies of the two sets of interacting species are supplied,
two phyogenetic signal strength parameters (d1 and d2), one for each species set, based on an Ornstein-Uhlenbeck model of
evolution with stabilizing selection are estimated. Values of d=1 indicate no stabilizing selection and correspond to the Brownian motion model of
evolution; 0<d<1 represents stabilizing selection; d=0 depicts the absence of phylogenetic correlation (i.e., a star phylogeny); and d>1 corresponds
to disruptive selection where phylogenetic signal is amplified. Confidence intervals for these and the other parameters can be estimated with
bootstrapping.
The function pblmpredict predicts the associations of novel species following the methods given in appendix B of Ives and Godfray (2006).
MSE |
total, full (each d estimated), star (d=0), and base (d=1) mean squared errors |
signal.strength |
two estimates of phylogenetic signal strength |
coefficients |
estimated intercept and covariate coefficients with approximate 95 percent CIs for the three model types (full, star, base) |
CI.boot |
95 percent CIs for all parameters |
variates |
matrix of model variates (can be used for plotting) |
residuals |
matrix of residuals from the three models (full, star and base) |
predicted |
predicted associations |
bootvalues |
matrix of parameters estimated from the |
phylocovs |
phylogenetic covariance matricies scaled by the estimated |
cors.1 |
correlations among predicted and observed associations for species of |
cors.2 |
correlations among predicted and observed associations for species of |
pred.novels1 |
predicted associations for the novel speices of |
pred.novels2 |
predicted associations for the novel speices of |
Covariates that apply to both species sets (e.g., sampling site) should be supplied in the covariate matrix of the set with the most species.
Bootstrapping CIs is slow due to the function optim used to estimate the model parameters. See appendix A in Ives and Godfray (2006)
for a discussion about this boostrapping procedure
If pblmpredict=TRUE the function does not first remove each species in turn when predicting the associations of the original species as
is done in Ives and Godfray (2006).
Matthew Helmus [email protected]
Ives A.R. & Godfray H.C. (2006) Phylogenetic analysis of trophic associations. The American Naturalist, 168, E1-E14
Blomberg S.P., Garland T.J. & Ives A.R. (2003) Testing for phylogenetic signal in comparative data: Behavioral traits are more labile. Evolution, 57, 717-745
#load example data from Ives & Godfray (2006) data(IvesGodfray) #net attack rate of parasitoid on host eq.4 in Ives and Godfray A<-(-1*log(1-IvesGodfray$interactions[,-28]/t(IvesGodfray$interactions[28]))) # Make tips of the phylogenetic trees contemporaneous by extending tips p<-dim(IvesGodfray$host)[1] q<-dim(IvesGodfray$parasitoid)[1] host.cov.scaled<-IvesGodfray$host para.cov.scaled<-IvesGodfray$parasitoid for (i in 1:p) { host.cov.scaled[i,i]<-max(host.cov.scaled) } for (i in 1:q) { para.cov.scaled[i,i]<-max(para.cov.scaled) } # scale covariance matrices (this reduces numerical problems caused by # determinants going to infinity or zero) host.cov.scaled<-host.cov.scaled/(det(as.matrix(host.cov.scaled))^(1/p)) para.cov.scaled<-para.cov.scaled/(det(as.matrix(para.cov.scaled))^(1/q)) pblm.A <- pblm(sqrt(A),tree1=host.cov.scaled,tree2=para.cov.scaled) pblm.A$signal.strength #compare to Ives and Godfray (2006) Table 1 Line 1 pblm.A$MSE#load example data from Ives & Godfray (2006) data(IvesGodfray) #net attack rate of parasitoid on host eq.4 in Ives and Godfray A<-(-1*log(1-IvesGodfray$interactions[,-28]/t(IvesGodfray$interactions[28]))) # Make tips of the phylogenetic trees contemporaneous by extending tips p<-dim(IvesGodfray$host)[1] q<-dim(IvesGodfray$parasitoid)[1] host.cov.scaled<-IvesGodfray$host para.cov.scaled<-IvesGodfray$parasitoid for (i in 1:p) { host.cov.scaled[i,i]<-max(host.cov.scaled) } for (i in 1:q) { para.cov.scaled[i,i]<-max(para.cov.scaled) } # scale covariance matrices (this reduces numerical problems caused by # determinants going to infinity or zero) host.cov.scaled<-host.cov.scaled/(det(as.matrix(host.cov.scaled))^(1/p)) para.cov.scaled<-para.cov.scaled/(det(as.matrix(para.cov.scaled))^(1/q)) pblm.A <- pblm(sqrt(A),tree1=host.cov.scaled,tree2=para.cov.scaled) pblm.A$signal.strength #compare to Ives and Godfray (2006) Table 1 Line 1 pblm.A$MSE
Pairwise dissimilarity in phylogenetic community composition that is partitioned into a nonphylogenetic and a phylogenetic component.
pcd(comm, tree, PSVmncd=NULL, PSVpool=NULL, reps=10^4)pcd(comm, tree, PSVmncd=NULL, PSVpool=NULL, reps=10^4)
comm |
Community data matrix |
tree |
Object of class phylo or a phylogenetic covariance matrix |
PSVmncd |
Vector of null mean conditional phylogenetic species variability (PSV) values |
PSVpool |
The standard, unconditional PSV calculated for the species pool |
reps |
The number of random draws from the species pool used to produce |
Phylogenetic community dissimilarity (PCD) is the pairwise differences between communities derived by asking how much of the variance
among species in the values of a hypothetical nonselected trait in one community can be predicted by the known trait values of species in another community.
PCD is partitioned into a nonphylogenetic component that reflects shared species between communities (PCDc)
and a phylogenetic component that reflects the evolutionary relationships among nonshared species (PCDp). In order to compare communities that vary
in species richness, the metric is standardized under the assumption that the species in communities are selected at random from the species pool. The
analyses here define the species pool as the list of all species in the set of communities in comm, but the species pool can be defined under
any hypothesis of community assembly either by manipulating the code or inputting a user defined PSVmncd and PSVpool.
The function returns a list with items:
PCD |
A square matrix of PCD values |
PCDc |
A square matrix of PCDc values |
PCDp |
A square matrix of PCDp values |
PSVmncd |
A vector of null mean conditional PSV values used to calculate PCD |
PSVpool |
The unconditional PSV of the species pool used to calculate PCD |
The sampling procedure used to standardize PCD and produce PSVmncd and PSVpool can be slow.
Anthony Ives <[email protected]> and Matthew Helmus <[email protected]>
Ives A.R. & Helmus M.R. (2010). Phylogenetic metrics of community similarity. The American Naturalist, 176, E128-E142.
data(phylocom) pcd(phylocom$sample, phylocom$phylo)data(phylocom) pcd(phylocom$sample, phylocom$phylo)
Calculate the sum of the total phylogenetic branch length for one or multiple samples.
pd(samp, tree, include.root=TRUE)pd(samp, tree, include.root=TRUE)
samp |
Community data matrix |
tree |
A phylo tree object |
include.root |
Should the root node be included in all PD calculations (default = TRUE) |
Returns a dataframe of the PD and species richness (SR) values for all samples
If the root is to be included in all calculations (include.root=TRUE), the PD of all samples will include the branch length connecting taxa in those samples and the root node of the supplied tree. The root of the supplied tree may not be spanned by any taxa in the sample. If you want the root of your tree to correspond to the most recent ancestor of the taxa actually present in your sample, you should prune the tree before running pd:
prunedTree <- prune.sample(sample,tree)
The data sets need not be species-community data sets but may be any sample data set with an associated phylogeny. PD is not statistically independent of species richness, it positively correlates with species richness across samples. The function ses.pd compares observed PD to the values expected under various randomizations and allows a way to standardize for unequal richness across samples.
If the root is to be included in all calculations of PD (include.root=TRUE), the tree must be rooted. Single-species samples will be assigned a PD value equal to the distance from the root to the present.
If the root is not included in all calculations by default (include.root=FALSE), the tree need not rooted, but in the case of single-species samples the PD will be equal to NA and a warning will be issued.
Matthew Helmus [email protected], Jonathan Davies [email protected], Steven Kembel [email protected]
Faith D.P. (1992) Conservation evaluation and phylogenetic diversity. Biological Conservation, 61, 1-10.
data(phylocom) pd(phylocom$sample, phylocom$phylo)data(phylocom) pd(phylocom$sample, phylocom$phylo)
Uses phylogenetic ancestral state reconstruction to estimate trait values for unobserved taxa.
phyEstimate(phy, trait, method = "pic", ...)phyEstimate(phy, trait, method = "pic", ...)
phy |
phylo object |
trait |
vector or data.frame containing trait values |
method |
ancestral state estimation method used by |
... |
Additional arguments passed to |
best.state |
estimate best-supported trait state for discrete variables? (default=TRUE) |
cutoff |
support cutoff required to declare a best.state |
These functions use phylogenetic ancestral state estimation to infer trait
values for novel taxa on a phylogenetic tree, for continuous
(phyEstimate) and discrete (phyEstimateDisc) traits.
The required input is a phylogenetic tree object plus a vector or data.frame containing estimated trait values for a subset of the taxa in the phylogenetic tree. Trait values for taxa that are present in the tree but not the trait data will be estimated using ancestral state estimation (Garland and Ives 2000). Briefly, for each taxon present in the tree but not the trait data, the phylogeny is rerooted at the most recent common ancestor of the novel taxon and the rest of the phylogeny, and the trait value of the novel taxon is estimated from the reconstructed trait value at the root of the rerooted phylogeny.
For phyEstimateDisc, the state with the highest support will be
reported if argument best.state=TRUE. If the best-supported state's
support is less than the specified cutoff, no best state is reported
and a NA value will be returned.
phyEstimate produces a data frame with columns:
est |
Estimated trait value |
se |
Standard error of estimated trait value |
phyEstimateDisc produces a data frame with columns:
states 1..N |
A column with statistical support is produced for each discrete trait state |
estimated.state |
If best.state=TRUE, a column with the state with the highest support |
estimated.state.support |
Statistical support for the state with the highest support |
Steven Kembel [email protected]
T. Garland Jr., and A.R. Ives. 2000. Using the past to predict the present: confidence intervals for regression equations in phylogenetic comparative methods. American Naturalist 155:346364.
S.W. Kembel, M. Wu, J.A. Eisen, and J.L. Green. 2012. Incorporating 16S gene copy number information improves estimates of microbial diversity and abundance. PLoS Computational Biology 8(10):e1002743.
#generate random phylogeny randtree <- rcoal(50) #simulate trait evolution for a subset of taxa on phylogeny randtraits <- sample(rTraitCont(randtree, sigma=10, root.value=100), 40) #estimate trait values for "missing" taxa using PIC method phyEstimate(randtree, randtraits, method="pic")#generate random phylogeny randtree <- rcoal(50) #simulate trait evolution for a subset of taxa on phylogeny randtraits <- sample(rTraitCont(randtree, sigma=10, root.value=100), 40) #estimate trait values for "missing" taxa using PIC method phyEstimate(randtree, randtraits, method="pic")
Phylogeny, community and trait data from the Phylocom 4.0 distribution
data(phylocom)data(phylocom)
A list with three elements:
phylo Phylogenetic tree (an object of class phylo)
sample Community data (a data.frame with samples in rows and species in columns
traits Trait data (a data.frame with species in rows and traits in columns
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
Calculates K statistic of phylogenetic signal as well as P-value based on variance of phylogenetically independent contrasts relative to tip shuffling randomization.
phylosignal(x, phy, reps = 999, checkdata=TRUE, ...)phylosignal(x, phy, reps = 999, checkdata=TRUE, ...)
x |
Trait vector (same order as phy\$tip.label) |
phy |
phylo object |
reps |
Number of randomizations |
checkdata |
Check for match between trait and phylogeny taxa labels using |
... |
Additional arguments passed to pic |
Data frame with columns:
K |
K statistic |
PIC.variance |
Mean observed PIC variance |
PIC.variance.P |
P-value of observed vs. random variance of PICs |
PIC.variance.z |
Z-score of observed vs. random variance of PICs |
Steven Kembel <[email protected]>
Blomberg, S. P., and T. Garland, Jr. 2002. Tempo and mode in evolution: phylogenetic inertia, adaptation and comparative methods. Journal of Evolutionary Biology 15:899-910.
Blomberg, S. P., T. Garland, Jr., and A. R. Ives. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717-745.
randtree <- rcoal(20) randtraits <- rTraitCont(randtree) phylosignal(randtraits[randtree$tip.label],randtree)randtree <- rcoal(20) randtraits <- rTraitCont(randtree) phylosignal(randtraits[randtree$tip.label],randtree)
Fraction of branch-length shared between two communities
phylosor(samp, tree)phylosor(samp, tree)
samp |
Community data matrix |
tree |
Object of class phylo - a rooted phylogeny |
A distance object of the PhyloSor index of similarity between communities, the fraction of PD (branch-length) shared between two samples
The phylosor of all samples will include the branch length connecting taxa in those samples and the root of the supplied tree. The root of the supplied tree may not be spanned by any taxa in the sample. If you want the root of your tree to correspond to the most recent ancestor of the taxa actually present in your sample, you should prune the tree before running phylosor:
prunedTree <- prune.sample(sample,tree)
The root of the supplied tree is included in calculations of PhyloSor. The supplied tree must be rooted. Single-species samples will be assigned a PD value equal to the distance from the root to the present.
Helene Morlon <[email protected]> and Steven Kembel <[email protected]>
Bryant, J.B., Lamanna, C., Morlon, H., Kerkhoff, A.J., Enquist, B.J., Green, J.L. 2008. Microbes on mountainsides: Contrasting elevational patterns of bacterial and plant diversity. Proceedings of the National Academy of Sciences 105 Supplement 1: 11505-11511
data(phylocom) phylosor(phylocom$sample, phylocom$phylo)data(phylocom) phylosor(phylocom$sample, phylocom$phylo)
PhyloSor values obtained by randomization for different choices of null models
phylosor.rnd(samp,tree, cstSor=TRUE, null.model=c("taxa.labels", "frequency","richness","independentswap","trialswap"), runs=999, iterations=1000)phylosor.rnd(samp,tree, cstSor=TRUE, null.model=c("taxa.labels", "frequency","richness","independentswap","trialswap"), runs=999, iterations=1000)
samp |
Community data matrix |
tree |
Object of class phylo - a rooted phylogeny |
cstSor |
TRUE if the Sorensen similarity should be kept constant across communities. FALSE otherwise |
null.model |
Null model to use (see Details section) |
runs |
Number of randomizations |
iterations |
Number of iterations to use for each randomization (for independent swap and trial null models) |
Currently implemented null models (arguments to null.model):
Shuffle community data matrix labels. Maintains species richness in each community and species shared between communities. Should be used with cstSor=TRUE
Randomize community data matrix abundances within species (maintains species occurence frequency). Does not maintain species richness in communities nor species shared between communities. Can only be used with cstSor=FALSE
With cstSor=TRUE: For each pair of community, maintains species richness in each community and species shared between communities. Sample in the species pool with equal probability; With cstSor=FALSE: Maintains species richness in each community, does not maintain species shared between communities. Sample in the species pool with equal probability
Randomize community data matrix with the independent swap algorithm (Gotelli 2000) maintaining species occurrence frequency and sample species richness. Can only be used with cstSor=FALSE
Randomize community data matrix with the trial-swap algorithm (Miklos & Podani 2004) maintaining species occurrence frequency and sample species richness. Can only be used with cstSor=FALSE
A list of length the number of runs. Each element of the list is a distance matrix containing the PhyloSor values of phylogenetic beta-diversity obtained by randomization
Helene Morlon <[email protected]> and Steven Kembel <[email protected]>
Bryant, J.B., Lamanna, C., Morlon, H., Kerkhoff, A.J., Enquist, B.J., Green, J.L. 2008. Microbes on mountainsides: Contrasting elevational patterns of bacterial and plant diversity. Proceedings of the National Academy of Sciences 105 Supplement 1: 11505-11511
data(phylocom) phylosor.rnd(phylocom$sample,phylocom$phylo,cstSor=TRUE,null.model="richness",runs=5)data(phylocom) phylosor.rnd(phylocom$sample,phylocom$phylo,cstSor=TRUE,null.model="richness",runs=5)
Randomize sample/community data matrices to create null distributions of given metrics
phylostruct(samp, tree, env=NULL, metric=c("psv","psr","pse","psc","sppregs"), null.model=c("frequency", "richness","independentswap","trialswap"), runs=100, it=1000, alpha=0.05, fam="binomial")phylostruct(samp, tree, env=NULL, metric=c("psv","psr","pse","psc","sppregs"), null.model=c("frequency", "richness","independentswap","trialswap"), runs=100, it=1000, alpha=0.05, fam="binomial")
samp |
community data matrix, species as columns, communities as rows |
tree |
phylo tree object or a phylogenetic covariance matrix |
env |
environmental data matrix |
metric |
if |
null.model |
permutation procedure used to create the null distribution, see |
runs |
the number of permutations to create the distribution, a rule of thumb is (number of communities)/alpha |
it |
the number of swaps for the independent and trial-swap null models, see |
alpha |
probability value to compare the observed mean/correlations to a null distribution |
fam |
as in |
The function creates null distributions for the psd set of metrics and for the correlations of sppregs from observed community data sets.
metric |
metric used |
null.model |
permutation used |
runs |
number of permutations |
it |
number of swaps if applicable |
obs |
observed mean value of a particular metric or the three observed correlations from |
mean.null |
mean(s) of the null distribution(s) |
quantiles.null |
quantiles of the null distribution(s) to compare to |
phylo.structure |
if |
nulls |
null values of the distribution(s) |
Matthew Helmus [email protected]
Helmus M.R., Bland T.J., Williams C.K. & Ives A.R. (2007a) Phylogenetic measures of biodiversity. American Naturalist, 169, E68-E83
Helmus M.R., Savage K., Diebel M.W., Maxted J.T. & Ives A.R. (2007b) Separating the determinants of phylogenetic community structure. Ecology Letters, 10, 917-925
Gotelli N.J. (2000) Null model analysis of species co-occurrence patterns. Ecology, 81, 2606-2621
Prune a phylogenetic tree to include only species present in a community data set or with non-missing trait data
prune.sample(samp, phylo) prune.missing(x, phylo)prune.sample(samp, phylo) prune.missing(x, phylo)
phylo |
phylo object |
samp |
Community data matrix |
x |
Vector of trait data |
Returns a pruned phylo object
Steven Kembel <[email protected]>
Calculate the bounded phylogenetic biodiversity metrics: phylogenetic species variability, richness, evenness and clustering for one or multiple samples.
psv(samp,tree,compute.var=TRUE,scale.vcv=TRUE) psr(samp,tree,compute.var=TRUE,scale.vcv=TRUE) pse(samp,tree,scale.vcv=TRUE) psc(samp,tree,scale.vcv=TRUE) psd(samp,tree,compute.var=TRUE,scale.vcv=TRUE) psv.spp(samp,tree)psv(samp,tree,compute.var=TRUE,scale.vcv=TRUE) psr(samp,tree,compute.var=TRUE,scale.vcv=TRUE) pse(samp,tree,scale.vcv=TRUE) psc(samp,tree,scale.vcv=TRUE) psd(samp,tree,compute.var=TRUE,scale.vcv=TRUE) psv.spp(samp,tree)
samp |
Community data matrix |
tree |
A phylo tree object or a phylogenetic covariance matrix |
compute.var |
Computes the expected variances for PSV and PSR for each community |
scale.vcv |
Scale the phylogenetic covariance matrix to bound the metric between 0 and 1 |
Phylogenetic species variability (PSV) quantifies how phylogenetic relatedness decreases the variance of a hypothetical unselected/neutral trait
shared by all species in a community. The expected value of PSV is statistically independent of species richness, is one when all species in a sample
are unrelated (i.e., a star phylogeny) and approaches zero as species become more related. PSV is directly related to mean phylogenetic distance, except
except calculated on a scaled phylogenetic covariance matrix. The expected variance around PSV for any sample of a particular species richness can be approximated.
To address how individual species contribute to the mean PSV of a data set, the function psv.spp gives signed proportions of the total deviation
from the mean PSV that occurs when all species are removed from the data set one at a time. The absolute values of these “species effects”
tend to positively correlate with species prevalence.
Phylogenetic species richness (PSR) is the number of species in a sample multiplied by PSV. It can be considered the species richness of a sample
after discounting by species relatedness. The value is maximum at the species richness of the sample, and decreases towards zero as relatedness
increases. The expected variance around PSR for any sample of a particular species richness can be approximated.
Phylogenetic species evenness (PSE) is the metric PSV modified to incorporate relative species abundances. The maximum attainable value of PSE (i.e., 1)
occurs only if species abundances are equal and species phylogeny is a star. PSE essentially grafts each individual of a species onto the tip of the
phylogeny of its species with branch lengths of zero.
Phylogenetic species clustering (PSC) is a metric of the branch tip clustering of species across a sample's phylogeny. As PSC increases to 1, species
are less related to one another the tips of the phylogeny. PSC is directly related to mean nearest neighbor distance.
Returns a dataframe of the respective phylogenetic species diversity metric values
These metrics are bounded either between zero and one (PSV, PSE, PSC) or zero and species richness (PSR); but the metrics asymptotically approach zero as relatedness increases. Zero can be assigned to communities with less than two species, but conclusions drawn from assigning communities zero values need be carefully explored for any data set. The data sets need not be species-community data sets but may be any sample data set with an associated phylogeny.
Matthew Helmus [email protected]
Helmus M.R., Bland T.J., Williams C.K. & Ives A.R. (2007) Phylogenetic measures of biodiversity. American Naturalist, 169, E68-E83
data(phylocom) psd(phylocom$sample, phylocom$phylo)data(phylocom) psd(phylocom$sample, phylocom$phylo)
Various null models for randomizing community data matrices
randomizeMatrix(samp, null.model = c("frequency", "richness", "independentswap", "trialswap"), iterations = 1000)randomizeMatrix(samp, null.model = c("frequency", "richness", "independentswap", "trialswap"), iterations = 1000)
samp |
Community data matrix |
null.model |
Null model to use (see Details section for description) |
iterations |
Number of independent or trial-swaps to perform |
Currently implemented null models (arguments to null.model):
Randomize community data matrix abundances within species (maintains species occurence frequency)
Randomize community data matrix abundances within samples (maintains sample species richness)
Randomize community data matrix with the independent swap algorithm (Gotelli 2000) maintaining species occurrence frequency and sample species richness
Randomize community data matrix with the trial-swap algorithm (Miklos & Podani 2004) maintaining species occurrence frequency and sample species richness
Randomized community data matrix
Steven Kembel <[email protected]>
Gotelli, N.J. 2000. Null model analysis of species co-occurrence patterns. Ecology 81: 2606-2621
Miklos I. & Podani J. 2004. Randomization of presence-absence matrices: Comments and new algorithms. Ecology 85: 86-92.
data(phylocom) randomizeMatrix(phylocom$sample, null.model="richness")data(phylocom) randomizeMatrix(phylocom$sample, null.model="richness")
Calculates Rao's quadratic entropy, a measure of within- and among-community diversity taking species dissimilarities into account
raoD(comm, phy=NULL)raoD(comm, phy=NULL)
comm |
Community data matrix |
phy |
Object of class phylo - an ultrametric phylogenetic tree (optional) |
Rao's quadratic entropy (Rao 1982) is a measure of diversity in ecological communities that can optionally take species differences (e.g. phylogenetic dissimilarity) into account. This method is conceptually similar to analyses of genetic diversity among populations (Nei 1973), but instead of diversity of alleles among populations, it measures diversity of species among communities.
If no phylogeny is supplied, Dkk is equivalent to Simpson's diversity (probability that two individuals drawn from a community are from different taxa), Dkl is a beta-diversity equivalent of Simpson's diversity (probability that individuals drawn from each of two communities belong to different taxa), and H is Dkl standardized to account for within-community diversity.
If an ultrametric phylogeny is supplied, Dkk is equivalent to the mean pairwise phylogenetic distance (distance to MRCA) between two individuals drawn from a community, Dkl is the mean pairwise phylogenetic distance between individuals drawn from each of two communities, and H is Dkl standardized to account for within-community diversity.
D[kl] = sum(t[ij] * x[ki] * x[lj])
where x[ki] is the relative abundance of taxon i in community k and t[ij] is a matrix of weights for all pairs of taxa i,j. Without a phylogeny, when i=j, t[ij]=0, otherwise t[ij]=1. With a phylogeny, t[ij] is the phylogenetic distance to MRCA for taxa i,j.
H[kl] = D[kl] - (D[kk] + D[ll])/2
Alpha, beta and total measure the average diversity within, among, and across all communities based on Dkk and H values taking variation in number of individuals per community into account. A Fst-like measure is calculated by dividing beta by the total diversity across all samples.
A list of results
Dkk |
Within-community diversity |
Dkl |
Among-community diversity |
H |
Among-community diversities excluding within-community diversity |
total |
Total diversity across all samples |
alpha |
Alpha diversity - average within-community diversity |
beta |
Beta diversity - average among-community diversity |
Fst |
Beta diversity / total diversity |
Alpha, beta, and total diversity components and Fst should not be interpreted as a measure of relative differentiation among versus within communities. See Jost (2007) for a detailed description of this problem. Hardy and Jost (2008) suggest Fst can be interpreted as 'local species identity excess' or 'local phylogenetic similarity excess' rather than as a measure of among-community differentiation.
Steven Kembel <[email protected]>
Hardy, O.J., and Jost. L. 2008. Interpreting and estimating measures of community phylogenetic structuring. J. Ecol. 96:849-852.
Jost, L. 2007. Partitioning diversity into independent alpha and beta components. Ecology 88: 24272439.
Nei, M. 1973. Analysis of gene diversity in sub-divided populations. Proceedings of the National Academy of Sciences of the USA 70:3321-3323.
Rao, C.R. 1982. Diversity and dissimilarity coefficients: a unified approach. Theoretical Population Biology 21:2443.
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
data(phylocom) raoD(phylocom$sample) raoD(phylocom$sample, phylocom$phylo)data(phylocom) raoD(phylocom$sample) raoD(phylocom$sample, phylocom$phylo)
Reads a Phylocom sample file and converts to a community data matrix
readsample(filename = "")readsample(filename = "")
filename |
Phylocom sample file path |
Community data matrix
Steven Kembel <skembel> and Cam Webb <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
Convert a Phylocom database-format sample to community data matrix.
sample2matrix(x)sample2matrix(x)
x |
Phylocom sample formatted data frame, a data frame with three columns:
|
Steven Kembel <[email protected]> and Cam Webb <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
Standardized effect size of mean nearest taxon distances in communities. When used with a phylogenetic distance matrix, equivalent to -1 times the Nearest Taxon Index (NTI).
ses.mntd(samp, dis, null.model = c("taxa.labels", "richness", "frequency", "sample.pool", "phylogeny.pool", "independentswap", "trialswap"), abundance.weighted=FALSE, runs = 999, iterations = 1000)ses.mntd(samp, dis, null.model = c("taxa.labels", "richness", "frequency", "sample.pool", "phylogeny.pool", "independentswap", "trialswap"), abundance.weighted=FALSE, runs = 999, iterations = 1000)
samp |
Community data matrix |
dis |
Distance matrix (generally a phylogenetic distance matrix) |
null.model |
Null model to use (see Details section for description) |
abundance.weighted |
Should mean nearest taxon distances for each species be weighted by species abundance? (default = FALSE) |
runs |
Number of randomizations |
iterations |
Number of iterations to use for each randomization (for independent swap and trial null models) |
The metric used by this function has also been referred to as MNND (mean nearest neighbour distance), and the function was named ses.mnnd in picante versions < 0.7.
Currently implemented null models (arguments to null.model):
Shuffle distance matrix labels (across all taxa included in distance matrix)
Randomize community data matrix abundances within samples (maintains sample species richness)
Randomize community data matrix abundances within species (maintains species occurence frequency)
Randomize community data matrix by drawing species from pool of species occurring in at least one community (sample pool) with equal probability
Randomize community data matrix by drawing species from pool of species occurring in the distance matrix (phylogeny pool) with equal probability
Randomize community data matrix with the independent swap algorithm (Gotelli 2000) maintaining species occurrence frequency and sample species richness
Randomize community data matrix with the trial-swap algorithm (Miklos & Podani 2004) maintaining species occurrence frequency and sample species richness
A data frame of results for each community
ntaxa |
Number of taxa in community |
mntd.obs |
Observed MNTD in community |
mntd.rand.mean |
Mean MNTD in null communities |
mntd.rand.sd |
Standard deviation of MNTD in null communities |
mntd.obs.rank |
Rank of observed MNTD vs. null communities |
mntd.obs.z |
Standardized effect size of MNTD vs. null communities (= (mntd.obs - mntd.rand.mean) / mntd.rand.sd, equivalent to -NTI) |
mntd.obs.p |
P-value (quantile) of observed MNTD vs. null communities (= mntd.obs.rank / runs + 1) |
runs |
Number of randomizations |
Steven Kembel <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
data(phylocom) ses.mntd(phylocom$sample, cophenetic(phylocom$phylo),null.model="taxa.labels")data(phylocom) ses.mntd(phylocom$sample, cophenetic(phylocom$phylo),null.model="taxa.labels")
Standardized effect size of mean pairwise distances in communities. When used with a phylogenetic distance matrix, equivalent to -1 times the Nearest Relative Index (NRI).
ses.mpd(samp, dis, null.model = c("taxa.labels", "richness", "frequency", "sample.pool", "phylogeny.pool", "independentswap", "trialswap"), abundance.weighted = FALSE, runs = 999, iterations = 1000)ses.mpd(samp, dis, null.model = c("taxa.labels", "richness", "frequency", "sample.pool", "phylogeny.pool", "independentswap", "trialswap"), abundance.weighted = FALSE, runs = 999, iterations = 1000)
samp |
Community data matrix |
dis |
Distance matrix (generally a phylogenetic distance matrix) |
null.model |
Null model to use (see Details section for description) |
abundance.weighted |
Should mean nearest taxon distances for each species be weighted by species abundance? (default = FALSE) |
runs |
Number of randomizations |
iterations |
Number of iterations to use for each randomization (for independent swap and trial null models) |
Currently implemented null models (arguments to null.model):
Shuffle distance matrix labels (across all taxa included in distance matrix)
Randomize community data matrix abundances within samples (maintains sample species richness)
Randomize community data matrix abundances within species (maintains species occurence frequency)
Randomize community data matrix by drawing species from pool of species occurring in at least one community (sample pool) with equal probability
Randomize community data matrix by drawing species from pool of species occurring in the distance matrix (phylogeny pool) with equal probability
Randomize community data matrix with the independent swap algorithm (Gotelli 2000) maintaining species occurrence frequency and sample species richness
Randomize community data matrix with the trial-swap algorithm (Miklos & Podani 2004) maintaining species occurrence frequency and sample species richness
A data frame of results for each community
ntaxa |
Number of taxa in community |
mpd.obs |
Observed mpd in community |
mpd.rand.mean |
Mean mpd in null communities |
mpd.rand.sd |
Standard deviation of mpd in null communities |
mpd.obs.rank |
Rank of observed mpd vs. null communities |
mpd.obs.z |
Standardized effect size of mpd vs. null communities (= (mpd.obs - mpd.rand.mean) / mpd.rand.sd, equivalent to -NRI) |
mpd.obs.p |
P-value (quantile) of observed mpd vs. null communities (= mpd.obs.rank / runs + 1) |
runs |
Number of randomizations |
Steven Kembel <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
data(phylocom) ses.mpd(phylocom$sample, cophenetic(phylocom$phylo),null.model="taxa.labels")data(phylocom) ses.mpd(phylocom$sample, cophenetic(phylocom$phylo),null.model="taxa.labels")
Standardized effect size of phylogenetic diversity (Faith's PD) in communities.
ses.pd(samp, tree, null.model = c("taxa.labels", "richness", "frequency", "sample.pool", "phylogeny.pool", "independentswap", "trialswap"), runs = 999, iterations = 1000, include.root=TRUE)ses.pd(samp, tree, null.model = c("taxa.labels", "richness", "frequency", "sample.pool", "phylogeny.pool", "independentswap", "trialswap"), runs = 999, iterations = 1000, include.root=TRUE)
samp |
Community data matrix |
tree |
Phylogeny (phylo object) |
null.model |
Null model to use (see Details section for description) |
runs |
Number of randomizations |
iterations |
Number of iterations to use for each randomization (for independent swap and trial null models) |
include.root |
Include distance to root node in calculation of PD (see documentation in |
Currently implemented null models (arguments to null.model):
Shuffle taxa labels across tips of phylogeny (across all taxa included in phylogeny)
Randomize community data matrix abundances within samples (maintains sample species richness)
Randomize community data matrix abundances within species (maintains species occurence frequency)
Randomize community data matrix by drawing species from pool of species occurring in at least one community (sample pool) with equal probability
Randomize community data matrix by drawing species from pool of species occurring in the phylogeny (phylogeny pool) with equal probability
Randomize community data matrix with the independent swap algorithm (Gotelli 2000) maintaining species occurrence frequency and sample species richness
Randomize community data matrix with the trial-swap algorithm (Miklos & Podani 2004) maintaining species occurrence frequency and sample species richness
A data frame of results for each community
ntaxa |
Number of taxa in community |
pd.obs |
Observed PD in community |
pd.rand.mean |
Mean PD in null communities |
pd.rand.sd |
Standard deviation of PD in null communities |
pd.obs.rank |
Rank of observed PD vs. null communities |
pd.obs.z |
Standardized effect size of PD vs. null communities (= (pd.obs - pd.rand.mean) / pd.rand.sd) |
pd.obs.p |
P-value (quantile) of observed PD vs. null communities (= mpd.obs.rank / runs + 1) |
runs |
Number of randomizations |
Steven Kembel <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
Proches, S., Wilson, J.R.U. and Cowling, R.M. 2006. How much evolutionary history in a 10 x 10m plot? Proceedings of Royal Society of London B, Biological Sciences 273:1143-1148.
data(phylocom) ses.pd(phylocom$sample, phylocom$phylo, null.model="taxa.labels", runs=99)data(phylocom) ses.pd(phylocom$sample, phylocom$phylo, null.model="taxa.labels", runs=99)
Finds a sample-based rarefaction curve for phylogentic species richness for a set of samples.
specaccum.psr(samp, tree, permutations = 100, method = "random", ...)specaccum.psr(samp, tree, permutations = 100, method = "random", ...)
samp |
Community data matrix |
tree |
A phylo tree object or a phylogenetic covariance matrix |
permutations |
Number of permutations with method |
method |
Species accumulation method, currently only |
... |
Other parameters to functions |
The function returns an object of class "specaccum" with items:
call |
Function call. |
method |
Accumulator method. |
sites |
Number of sites/samples. |
richness |
The mean phylogenetic species richness corresponding to number of sites/samples. |
sd |
The standard deviation of phylogenetic apecies accumulation curve (or its standard error) estimated from permutations in |
perm |
Permutation results with |
Matthew Helmus [email protected] based on the vegan package specaccum function by Roeland Kindt and Jari Oksanen.
Gotelli N.J. & Colwell R.K. (2001) Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters, 4, 379-391
Helmus M.R., Bland T.J., Williams C.K. & Ives A.R. (2007) Phylogenetic measures of biodiversity. American Naturalist, 169, E68-E83
data(phylocom) accum.sr<-specaccum(phylocom$sample, permutations = 100, method = "random") plot(accum.sr, col="blue") points(accum.sr$sites, accum.sr$richness, pch=19, col="blue") accum.psr<-specaccum.psr(phylocom$sample, phylocom$phylo, permutations = 100, method = "random") plot(accum.psr, add=TRUE, col = "red") points(accum.psr$sites, accum.psr$richness, pch=19, col="red") legend(5,5,legend=c("SR","PSR"),pch=c(19,19),col=c("blue","red"))data(phylocom) accum.sr<-specaccum(phylocom$sample, permutations = 100, method = "random") plot(accum.sr, col="blue") points(accum.sr$sites, accum.sr$richness, pch=19, col="blue") accum.psr<-specaccum.psr(phylocom$sample, phylocom$phylo, permutations = 100, method = "random") plot(accum.psr, add=TRUE, col = "red") points(accum.psr$sites, accum.psr$richness, pch=19, col="red") legend(5,5,legend=c("SR","PSR"),pch=c(19,19),col=c("blue","red"))
Compute interspecific distances based on patterns of species co-occurrence in communities.
species.dist(x, metric = c("cij", "jaccard", "checkerboard", "doij"))species.dist(x, metric = c("cij", "jaccard", "checkerboard", "doij"))
x |
Community data matrix |
metric |
Co-occurrence metric to use (see Details section for description) |
Currently implemented co-occurrence measures (arguments to metric):
Schoener's index of co-occurrence
Jaccard index of co-occurrence
Checkerboard index of co-occurrence
DOij index of co-occurrence
A dist object with co-occurrences among all species pairs
Steven Kembel <[email protected]>
Hardy, O.J. 2008. Testing the spatial phylogenetic structure of local communities: statistical performances of different null models and test statistics on a locally neutral community. Journal of Ecology 96:914-926.
Fit regressions on species abundance or presence/absence across communities and calculate phylogenetic correlations
sppregs(samp, env, tree=NULL, fam="gaussian") sppregs.plot(sppreg, rows=c(1,3), cex.mag=1, x.label="phylogenetic correlations", y.label=c("occurrence correlations w/ env", "occurrence correlations wo/ env", "change in correlations"))sppregs(samp, env, tree=NULL, fam="gaussian") sppregs.plot(sppreg, rows=c(1,3), cex.mag=1, x.label="phylogenetic correlations", y.label=c("occurrence correlations w/ env", "occurrence correlations wo/ env", "change in correlations"))
samp |
community data matrix, species as columns, communities as rows |
env |
environmental data matrix |
tree |
phylo tree object or a phylogenetic covariance matrix |
fam |
with |
sppreg |
object from function |
rows |
|
cex.mag |
value for |
x.label |
x axis labels |
y.label |
y axis labels |
For each species in samp, the function fits regressions of species presence/absence or abundances
on the environmental variables supplied in env; and calculates the (n^2-n)/2 pairwise species correlations
between the residuals of these fits and pairwise species phylogenetic correlations. The residuals can be
thought of as the presence/absence of species across sites/communities after accounting for how species respond
to environmental variation across sites. Each set of coefficients can be tested for phylogenetic signal with, for example,
the function phylosignal.
The function sppregs.plot produces a set of three plots of the correlations of pairwise species phylogenetic correlations versus:
the observed pairwise correlations of species across communities, the residual correlations, and the pairwise differences between (i.e., the
change in species co-occurrence once the environmental variables are taken into account). The significance of these correlations can be tested
via permutation with the function phylostruct.
family |
the regression error distribution |
residuals |
the residuals from each species regression |
coefficients |
the estimated coefficients from each species regression |
std.errors |
the standard errors of the coefficients |
correlations |
correlations of pairwise species phylogenetic correlations between: the observed pairwise correlations of species across communities, the residual correlations, and the pairwise differences between the two |
cors.pa |
the observed pairwise correlations of species across communities |
cors.resid |
the residual pairwise correlations of species across communities |
cors.phylo |
the phylogenetic pairwise correlations among species |
The function requires the library brglm to perform logistic regressions
Matthew Helmus [email protected]
Helmus M.R., Savage K., Diebel M.W., Maxted J.T. & Ives A.R. (2007) Separating the determinants of phylogenetic community structure. Ecology Letters, 10, 917-925
Taxic diversity: Vane-Wright et al., 1991 and May 1990 which accounts for polytomies by counting the number of branches descending from each node that lies on the path from a spp tip to the root (not just counting the number of nodes).
tax.distinctiveness(tree, type = c("Vane-Wright", "May"))tax.distinctiveness(tree, type = c("Vane-Wright", "May"))
tree |
an object of class phylo |
type |
specify "Vane-Wright" or "May" |
Karen Magnuson-Ford, Will Cornwell, Arne Mooers, Mark Vellend
Vane-Wright, R.I., Humphries, C.J. and Williams, P.H. (1991). What to protect? - Systematics and the agony of choice. Biological Conservation, 55, 235-254.
May, R.M. (1990). Taxonomy as destiny. Nature, 347, 129-130.
Mark Vellend, William K. Cornwell, Karen Magnuson-Ford, and Arne Mooers. In press. Measuring phylogenetic biodiversity In: Biological diversity: frontiers in measurement and assessment. Edited by Anne Magurran and Brian McGill.
Draws a phylogeny where x position of nodes and tips corresponds to value of a continuous trait variable, and y position corresponds to node depth (i.e. age).
traitgram(x, phy, xaxt = 's', underscore = FALSE, show.names = TRUE, show.xaxis.values = TRUE, method = c('ML','pic'), ...)traitgram(x, phy, xaxt = 's', underscore = FALSE, show.names = TRUE, show.xaxis.values = TRUE, method = c('ML','pic'), ...)
x |
Trait vector (same order as phy\$tip.label, or with taxon names in names) |
phy |
phylo object |
xaxt |
x axis default type |
underscore |
if FALSE remove underscore from taxonomic names |
show.names |
if TRUE show taxon names across tips of phylogeny |
show.xaxis.values |
if TRUE show values for trait on x=axis |
method |
method for calculation of internal trait values. 'ML' = maximum likelihood method; 'pic' = independent contrasts method. pic option can be used when ML fails to converge or otherwise seems to fail to correctly reconstruct ancestral values |
... |
Additional arguments passed to plot |
Plots a traitgram, no values returned.
David Ackerly <[email protected]>
Ackerly, D. D. 2009. Conservatism and diversification of plant functional traits: Evolutionary rates versus phylogenetic signal. Proceedings of the National Academy of Sciences USA 106:19699-19706. doi: 10.1073/pnas.0901635106.
Evans, M. E. K., S. A. Smith, R. S. Flynn, and M. J. Donoghue. 2009. Climate, Niche Evolution, and Diversification of the "bird-cage" Evening Primroses (Oenothera, Sections Anogra and Kleinia). American Naturalist 173:225-240.
randtree <- rcoal(20) randtraits <- rTraitCont(randtree) traitgram(randtraits,randtree) traitgram(randtraits,randtree,method='pic')randtree <- rcoal(20) randtraits <- rTraitCont(randtree) traitgram(randtraits,randtree) traitgram(randtraits,randtree,method='pic')
Calculates unweighted UniFrac, a phylogenetic beta diversity metric of the the unique (non-shared) fraction of total phylogenetic diversity (branch-length) between two communities.
unifrac(comm, tree)unifrac(comm, tree)
comm |
Community data matrix |
tree |
Object of class phylo - a rooted phylogeny |
A dist object of the unweighted UniFrac distances between communities (the unique (non-shared) fraction of total phylogenetic diversity (branch-length) between two communities).
The UniFrac distance between samples will include the branch length connecting taxa in those samples and the root of the supplied tree. The root of the supplied tree may not be spanned by any taxa in the sample. If you want the root of your tree to correspond to the most recent ancestor of the taxa actually present in your samples, you should prune the tree before running unifrac:
prunedTree <- prune.sample(sample,tree)
The supplied tree must be rooted. Single-species samples will be assigned a PD value equal to the distance from the root to the present.
Steven Kembel <[email protected]>
Lozupone, C., Hamady, M., and Knight, R. 2006. UniFrac - an online tool for comparing microbial community diversity in a phylogenetic context. BMC Bioinformatics 7:371.
data(phylocom) unifrac(phylocom$sample, phylocom$phylo)data(phylocom) unifrac(phylocom$sample, phylocom$phylo)
Picante utility functions for tree and data manipulation
df2vec(x, colID=1) internal2tips(phy, int.node, return.names = FALSE) node.age(phy) pic.variance(x, phy, scaled = TRUE) sortColumns(x) sortRows(x) taxaShuffle(x) tipShuffle(phy)df2vec(x, colID=1) internal2tips(phy, int.node, return.names = FALSE) node.age(phy) pic.variance(x, phy, scaled = TRUE) sortColumns(x) sortRows(x) taxaShuffle(x) tipShuffle(phy)
phy |
phylo object |
x |
A data.frame, matrix or dist object |
colID |
Numeric or character ID of column to include |
int.node |
internal node number |
return.names |
TRUE or FALSE |
scaled |
Scale contrasts by branch length |
Various utility functions for manipulating trees, data, etc.
df2vec |
A named vector |
internal2tips |
Vector of tips descended from a node |
node.age |
Phylo object with phylo\$ages vector of node ages corresponding to phylo\$edge |
pic.variance |
Variance of independent contrasts |
sortColumns |
A data.frame or matrix with columns sorted by name |
sortRows |
A data.frame or matrix with rows sorted by name |
taxaShuffle |
Matrix with taxa names shuffled |
tipShuffle |
Phylo object with taxa names shuffled |
Steven Kembel <[email protected]>, Peter Cowan <[email protected]>, David Ackerly <[email protected]>
Write a community data matrix to a Phylocom community sample file
writesample(community, filename = "")writesample(community, filename = "")
community |
Community data matrix |
filename |
Filename path |
Steven Kembel <[email protected]> and Cam Webb <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.
Write a Phylocom traits formatted file
writetraits(trt, file = "", bin = NULL, sigd = 3)writetraits(trt, file = "", bin = NULL, sigd = 3)
trt |
Data frame containing trait data |
file |
Filename path |
bin |
Vector index of trait columns to be treated as binary |
sigd |
Significant digits for output |
David Ackerly <[email protected]> and Steven Kembel <[email protected]>
Webb, C.O., Ackerly, D.D., and Kembel, S.W. 2008. Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Version 4.0.1. http://www.phylodiversity.net/phylocom/.