treespace implements new methods for the exploration and analysis of distributions of phylogenetic trees for a given set of taxa.
To install the development version from github:
The stable version can be installed from CRAN using:
Then, to load the package, use:
The main functions implemented in treespace are:
treespace
: explore landscapes of phylogenetic trees
treespaceServer
: open up an application in a web browser for an interactive exploration of the diversity in a set of trees
findGroves
: identify clusters of similar trees
plotGroves
: scatterplot of groups of trees, and plotGrovesD3
which enables interactive plotting based on d3.js
medTree
: find geometric median tree(s) to summarise a group of trees
Other functions are central to the computations of distances between trees:
treeVec
: characterise a tree by a vector
treeDist
: find the distance between two tree vectors
multiDist
: find the pairwise distances of a list of trees
refTreeDist
: find the distances of a list of trees from a reference tree
tipDiff
: for a pair of trees, list the tips with differing ancestry
plotTreeDiff
: plot a pair of trees, highlighting the tips with differing ancestry
Distributed datasets include:
woodmiceTrees
: illustrative set of 201 trees built using the neighbour-joining and bootstrapping example from the woodmice dataset in the ape documentation.
DengueTrees
: 500 trees sampled from a BEAST posterior set of trees from (Drummond and Rambaut, 2007)
DengueSeqs
: 17 dengue virus serotype 4 sequences from (Lanciotti et al., 1997), from which the DengueTrees
were inferred.
DengueBEASTMCC
: the maximum clade credibility (MCC) tree from the DengueTrees
.
We first load treespace, and the packages required for graphics:
The function treespace
defines typologies of phylogenetic trees using a two-step approach:
perform pairwise comparisons of trees using various (Euclidean) metrics; by default, the comparison uses the Kendall and Colijn metric (Kendall and Colijn, 2016) which is described in more detail below; other metrics rely on tip distances implemented in adephylo (Jombart et al., 2010) and phangorn (Schliep 2011).
use Metric Multidimensional Scaling (MDS, aka Principal Coordinates Analysis, PCoA) to summarise pairwise distances between the trees as well as possible into a few dimensions; the output of the MDS is typically visualised using scatterplots of the first few Principal Components (PCs); this step relies on the PCoA implemented in ade4 (Dray and Dufour, 2007).
The function treespace
performs both tasks, returning both the matrix of pairwise tree comparisons ($D
), and the PCoA ($pco
). This can be illustrated using randomly generated trees:
# generate list of trees
suppressWarnings(RNGversion("3.5.0"))
set.seed(1)
x <- rmtree(10, 20)
names(x) <- paste("tree", 1:10, sep = "")
# use treespace
res <- treespace(x, nf=3)
names(res)
## [1] "D" "pco"
## $D
## tree1 tree2 tree3 tree4 tree5 tree6 tree7 tree8 tree9
## tree2 26.00
## tree3 31.06 26.74
## tree4 42.85 42.12 44.44
## tree5 30.66 27.71 27.37 44.79
## tree6 36.50 31.18 30.18 41.81 31.59
## tree7 34.64 28.71 29.48 40.35 31.11 32.37
## tree8 28.97 26.29 24.45 43.74 23.47 30.41 29.00
## tree9 29.63 27.42 27.48 45.61 26.31 30.89 29.77 24.60
## tree10 34.87 30.00 29.44 44.97 34.06 31.05 34.41 31.54 32.59
##
## $pco
## Duality diagramm
## class: pco dudi
## $call: dudi.pco(d = D, scannf = is.null(nf), nf = nf)
##
## $nf: 3 axis-components saved
## $rank: 9
## eigen values: 142.1 76.52 62.69 49.88 41.07 ...
## vector length mode content
## 1 $cw 9 numeric column weights
## 2 $lw 10 numeric row weights
## 3 $eig 9 numeric eigen values
##
## data.frame nrow ncol content
## 1 $tab 10 9 modified array
## 2 $li 10 3 row coordinates
## 3 $l1 10 3 row normed scores
## 4 $co 9 3 column coordinates
## 5 $c1 9 3 column normed scores
## other elements: NULL
Pairwise tree distances can be visualised using adegraphics:
# table.value with some customization
table.value(res$D, nclass=5, method="color",
symbol="circle", col=redpal(5))
The best representation of these distances in a 2-dimensional space is given by the first 2 PCs of the MDS. These can be visualised using any scatter plotting tool; here we use the treespace function plotGroves
, based on the adegraphics function scatter
:
Alternatively, plotGrovesD3
creates interactive plots based on d3.js:
The functionality of treespace
can be further illustrated using ape’s dataset woodmouse, from which we built the 201 trees supplied in woodmiceTrees
using the neighbour-joining and bootstrapping example from the ape documentation.
data(woodmiceTrees)
wm.res <- treespace(woodmiceTrees,nf=3)
# PCs are stored in:
head(wm.res$pco$li)
## A1 A2 A3
## 1 -0.9949 -1.363 -0.7918
## 2 -0.6137 -1.014 -0.6798
## 3 2.6667 4.219 -2.9293
## 4 -13.6081 1.854 1.0947
## 5 2.1980 4.176 -3.1960
## 6 3.6013 4.865 2.9853
Packages such as adegraphics and ggplot2 can be used to make alternative plots, for example visualising the density of points within the space.
The treespace function multiDist
simply performs the pairwise comparison of trees and outputs a distance matrix. This function may be preferable for large datasets, and when principal co-ordinate analysis is not required. It includes an option to save memory at the expense of computation time.
Once a typology of trees has been derived using the approach described above, one may want to formally identify clusters of similar trees. One simple approach is:
select a few first PCs of the MDS (retaining signal but getting rid of random noise)
derive pairwise Euclidean distances between trees based on these PCs
use hierarchical clustering to obtain a dendrogram of these trees
cut the dendrogram to obtain clusters
In treespace, the function findGroves
implements this approach, offering various clustering options (see ?findGroves
). Here we supply the function with our treespace
output wm.res
since we have already calculated it, but it is also possible to skip the steps above and directly supply findGroves
with a multiPhylo list of trees.
## [1] "groups" "treespace"
Note that when the number of clusters (nclust
) is not provided, the function will display a dendrogram and ask for a cut-off height.
The results can be plotted directly using plotGrovesD3
(see ?plotGrovesD3
for options):
# alternative with improved legend and tooltip text, giving the tree numbers:
plotGrovesD3(wm.groves, tooltip_text=paste0("Tree ",1:201), legend_width=50, col_lab="Cluster")
# plot axes 2 and 3. This helps to show why, for example, clusters 2 and 4 have been identified as separate, despite them appearing to overlap when viewing axes 1 and 2.
plotGrovesD3(wm.groves, xax=2, yax=3, tooltip_text=paste0("Tree ",1:201), legend_width=50, col_lab="Cluster")
We can also plot in 3D:
# prepare a colour palette:
colours <- fac2col(wm.groves$groups, col.pal=funky)
plot3d(wm.groves$treespace$pco$li[,1],
wm.groves$treespace$pco$li[,2],
wm.groves$treespace$pco$li[,3],
col=colours, type="s", size=1.5,
xlab="", ylab="", zlab="")
You must enable Javascript to view this page properly.
treespaceServer
: a web application for treespaceThe functionalities of treespace
are also available via a user-friendly web interface, running locally on the default web browser. It can be started by simply typing treespaceServer()
. The interface allows you to import trees and run treespace
to view and explore the tree space in 2 or 3 dimensions. It is then straightforward to analyse the tree space by varying \(\lambda\), looking for clusters using findGroves
and saving results in various formats. Individual trees can be easily viewed, including median trees per cluster (see below). Pairs of trees can be viewed together with their tip-differences highlighted using the function plotTreeDiff
, and collections of trees can be seen together using densiTree
from the package phangorn. It is fully documented in the help tab.
When a set of trees have very similar structures, it makes sense to summarize them into a single ‘consensus’ tree. In treespace
, this is achieved by finding the median tree for a set of trees according to the Kendall and Colijn metric. That is, we find the tree which is closest to the centre of the set of trees in the tree landscape defined in treespace
. This procedure is implemented by the function medTree
:
# get first median tree
tre <- medTree(woodmiceTrees)$trees[[1]]
# plot tree
plot(tre,type="cladogram",edge.width=3, cex=0.8)