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#' Compute quartet distance between taxa
#'
#' Compute the Quartet Distance of \insertCite{Rho19;textual}{MSCquartets} from a table specifying a collection of quartets on
#' \code{n} taxa.
#'
#' @references
#' \insertRef{Rho19}{MSCquartets}
#'
#' @param dqt an (\code{n} choose 4) x \code{n} (or \code{n+1}) matrix of form output by \code{quartetTableDominant};
#' (Note: If present, the \code{n+1}th column of \code{dqt} is ignored.)
#'
#' @return a pairwise distance matrix on \code{n} taxa
#'
#' @seealso
#' \code{\link{quartetTableDominant}},
#' \code{\link{QDS}},
#' \code{\link{QDC}},
#' \code{\link{quartetWeightedDist}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' QT=quartetTable(gtrees,tnames[1:6])
#' RQT=quartetTableResolved(QT)
#' DQT=quartetTableDominant(RQT)
#' Dist=quartetDist(DQT)
#' tree=NJ(Dist)
#' write.tree(tree)
#' plot(tree)
#'
#' @export
quartetDist = function(dqt) {
Qcolnames = colnames(dqt)# get column names
n = length(Qcolnames) - 1# number of taxa
taxa = Qcolnames[1:n]# names of taxa
m = dim(dqt)[1]# number of quartets
D = matrix(0, n, n)# create distance matrix
colnames(D) = taxa
rownames(D) = taxa
for (i in 1:m) {
ones = which(dqt[i, ] == 1)
if (length(ones) == 2) {
# skip null quartets
nones = which(dqt[i, ] == -1)
D[ones, nones] = D[ones, nones] + 1 # increment 4 entries of D
D[nones, ones] = D[nones, ones] + 1 # and another 4
}
}
D = 2 * D + 2 * n - 4 # adjust to appropriate Q distance
for (i in 1:n) {
D[i, i] = 0 # and unadjust diagonal back to 0
}
return(D)
}
###################################################################
#' Compute Quartet Distance Supertree
#'
#' Apply the Quartet Distance Supertree method of \insertCite{Rho19;textual}{MSCquartets} to a table specifying a
#' collection of quartets on \code{n} taxa.
#'
#' @references
#' \insertRef{Rho19}{MSCquartets}
#'
#' @details This function is a wrapper which runs \code{quartetDist} and then builds a tree.
#'
#' @param dqt an (\code{n} choose 4) x \code{n} (or \code{n+1}) matrix of form output by \code{quartetTableDominant};
#' (Note: If present, the \code{n+1}th column of \code{dqt} is ignored)
#'
#' @param method tree building function (e.g., fastme.bal, nj)
#'
#' @return
#' an unrooted metric tree of type phylo. Edge lengths are not in interpretable units
#' @seealso
#' \code{\link{quartetTableDominant}},
#' \code{\link{quartetDist}},
#' \code{\link{QDC}},
#' \code{\link{WQDS}},
#' \code{\link{WQDC}},
#' \code{\link{WQDCrecursive}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' QT=quartetTable(gtrees,tnames[1:6])
#' RQT=quartetTableResolved(QT)
#' DQT=quartetTableDominant(RQT)
#' tree=QDS(DQT)
#' write.tree(tree)
#' plot(tree)
#'
#' @importFrom ape fastme.bal
#'
#' @export
QDS = function(dqt,
method = fastme.bal) {
D = quartetDist(dqt)
QDS = unroot(method(D))# build tree
return(QDS)
}
#################################################
#' Compute the Weighted Quartet Distance between taxa
#'
#' Compute the Weighted Quartet Distance between taxa of \insertCite{YR19;textual}{MSCquartets} from a table specifying a collection of quartets on
#' \code{n} taxa and the quartets' internal branch lengths.
#'
#' @references
#' \insertRef{YR19}{MSCquartets}
#'
#' @param dqt an (\code{n} choose 4) x (\code{n+1}) matrix of the form output by \code{quartetTableDominant}
#'
#' @return
#' a pairwise distance matrix on \code{n} taxa
#'
#' @seealso \code{\link{quartetTableDominant}},
#' \code{\link{WQDSAdjustLengths}},
#' \code{\link{WQDS}},
#' \code{\link{WQDC}},
#' \code{\link{WQDCrecursive}},
#' \code{\link{quartetWeightedDist}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' QT=quartetTable(gtrees,tnames[1:6])
#' RQT=quartetTableResolved(QT)
#' DQT=quartetTableDominant(RQT,bigweights="finite")
#' D=quartetWeightedDist(DQT)
#' tree=NJ(D)
#' stree=WQDSAdjustLengths(tree)
#' write.tree(stree)
#'
#' @export
quartetWeightedDist = function(dqt) {
Qcolnames = colnames(dqt)# get column names
n = length(Qcolnames) - 1# number of taxa
taxa = Qcolnames[1:n]# names of taxa
m = dim(dqt)[1]# number of quartets
w = dqt[, n + 1]# weights
D = matrix(2, n, n)# create distance matrix
colnames(D) = taxa
rownames(D) = taxa
for (i in 1:m) {
ones = which(dqt[i,] == 1)
if (length(ones) == 2) {
# skip null quartets
nones = which(dqt[i,] == -1)
D[ones, nones] = D[ones, nones] + w[i] # increment 4 entries of D
D[nones, ones] = D[nones, ones] + w[i] # and another 4
}
}
for (i in 1:n) {
D[i, i] = 0 # unadjust diagonal back to 0
}
return(D)
}
#################################################
#' Compute the Weighted Quartet Distance Supertree
#'
#' Apply the Weighted Quartet Distance Supertree method of \insertCite{YR19;textual}{MSCquartets} to
#' a collection of quartets on \code{n} taxa together with internal
#' quartet branch lengths, specified by a table.
#'
#' @references
#' \insertRef{YR19}{MSCquartets}
#'
#' @details This function is a wrapper which runs \code{quartetWeightedDist}, builds a tree, and then adjusts edge lengths
#' with \code{WQDSAdjustLengths}.
#'
#' @param dqt an (\code{n} choose 4) x \code{n+1}) matrix of form output by \code{quartetTableDominant}
#' @param method a distance-based tree building function (e.g., fastme.bal, NJ, etc.)
#'
#' @return an unrooted metric tree, of type phylo
#'
#' @seealso \code{\link{quartetTableDominant}},
#' \code{\link{quartetWeightedDist}},
#' \code{\link{WQDSAdjustLengths}},
#' \code{\link{WQDC}},
#' \code{\link{WQDCrecursive}},
#' \code{\link{QDS}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' QT=quartetTable(gtrees,tnames[1:6])
#' RQT=quartetTableResolved(QT)
#' DQT=quartetTableDominant(RQT,bigweights= "finite")
#' tree=WQDS(DQT)
#' write.tree(tree)
#' plot(tree)
#'
#' @importFrom ape fastme.bal
#' @export
WQDS = function(dqt,
method = fastme.bal) {
D = quartetWeightedDist(dqt)# compute distance
WQDS = unroot(method(D))# build tree
WQDS=WQDSAdjustLengths(WQDS)
return(WQDS)
}
#################################################
#' Adjust edge lengths on tree built from Weighted Quartet distance
#' to estimate metric tree
#'
#' Modify edge lengths of a tree built from a distance table produced by \code{quartetWeightedDist},
#' to remove scaling factors related to the size of the split associated to the edge.
#'
#' @details As explained by \insertCite{YR19;textual}{MSCquartets}, a metric tree produced from
#' the weighted quartet distance has edge lengths
#' inflated by a factor dependent on the associated split size. Removing these
#' factors
#' yields a consistent estimate of the metric species tree displaying the weighted
#' quartets, if such a tree exists.
#'
#' This function should not be used on trees output from \code{WQDS}, \code{WQDC},
#' or \code{WQDCrecursive}, as
#' their edges are already adjusted. It can be used on trees built from the distance
#' computed by \code{quartetWeightedDist}.
#'
#' @references
#' \insertRef{YR19}{MSCquartets}
#'
#' @param tree an unrooted metric tree, of type phylo
#'
#' @return an unrooted metric tree, of type phylo
#'
#' @seealso
#' \code{\link{WQDS}},
#' \code{\link{WQDC}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' QT=quartetTable(gtrees,tnames[1:6])
#' RQT=quartetTableResolved(QT)
#' DQT=quartetTableDominant(RQT,bigweights="finite")
#' D=quartetWeightedDist(DQT)
#' tree=NJ(D)
#' write.tree(tree)
#' plot(tree)
#' stree=WQDSAdjustLengths(tree)
#' write.tree(stree)
#' plot(stree)
#'
#' @importFrom ape is.rooted unroot root
#' @importFrom phangorn Descendants
#' @export
WQDSAdjustLengths = function(tree) {
if (is.rooted(tree) == TRUE) {
message("Tree is rooted; unrooting it.")
tree = unroot(tree)
}
ntree = root(tree, outgroup = tree$tip.label[1])
ntaxa = length(ntree$tip.label)
nedges = dim(ntree$edge)[1]
for (i in 1:nedges) {
node = ntree$edge[i, 2] # get child of edge
ndesc = length(Descendants(ntree, node, "tips")[[1]])# count descendents, using phangorn function
if (ndesc == 1) {
ntree$edge.length[i] = 1
}
else if (ndesc == ntaxa - 1) {
ntree$edge.length[i] = 0
}
else {
ntree$edge.length[i] = ntree$edge.length[i] / ((ndesc - 1) * (ntaxa - ndesc - 1)) # adjust edge length
}
}
WQDSAdjustLengths = unroot(ntree)
return(WQDSAdjustLengths)
}
#################################################
#' Compute Quartet Distance Consensus tree from gene tree data
#'
#' Compute the Quartet Distance Consensus \insertCite{Rho19}{MSCquartets} estimate of an unrooted
#' topological species tree from gene tree data.
#'
#' @details This function is a wrapper which performs the steps of reading in a collection
#' of gene trees, tallying quartets, computing the quartet distance between taxa, building
#' a tree which consistently estimates the unrooted species tree topology under the MSC, and then possibly estimating edge
#' lengths using the "simpleML" method.
#'
#' @references
#' \insertRef{Rho19}{MSCquartets}
#'
#' @param genetreedata gene tree data that may be supplied in any of 3 forms:
#' \enumerate{
#' \item a character string giving the name of a file containing gene trees in Newick,
#' \item a multiPhylo object containing gene trees, or
#' \item a resolved quartet table, such as produced by \code{quartetTableResolved}
#' }
#' @param taxanames if \code{genetreedata} is a file or a multiPhylo object, a vector of a subset
#' of the taxa names on the gene trees
#' to be analyzed, if \code{NULL} all taxa on the first gene tree are used; if \code{genetreedata}
#' is a quartet table, this argument is ignored and all taxa in the table are used
#' @param method a distance-based tree building function, such as \code{fastme.bal} or \code{nj}
#' @param omit \code{TRUE} ignores unresolved quartets; \code{FALSE} treats them as 1/3 of each resolution;
#' ignored if \code{genetreedata} is supplied as a quartet table
#' @param metric if \code{FALSE} return topological tree; if \code{TRUE} return metric tree with
#' internal edge lengths estimated by \code{estimateEdgeLengths} with \code{lambda=0}, and terminal branches of length 1
#'
#'
#' @return an unrooted tree of type phylo
#'
#' @seealso \code{\link{quartetTable}},
#' \code{\link{quartetTableResolved}},
#' \code{\link{quartetTableDominant}},
#' \code{\link{quartetDist}},
#' \code{\link{QDS}},
#' \code{\link{WQDC}},
#' \code{\link{WQDCrecursive}}
#' \code{\link{estimateEdgeLengths}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' stree=QDC(gtrees,tnames[1:6])
#' write.tree(stree)
#' plot(stree)
#' streeMetric=QDC(gtrees, tnames[1:6],metric=TRUE)
#' write.tree(streeMetric)
#' plot(streeMetric)
#'
#' @export
QDC = function(genetreedata,
taxanames = NULL,
method=fastme.bal,
omit = FALSE,
metric=FALSE) {
if ("matrix" %in% class(genetreedata)) {
RQT = genetreedata
if (!is.null(taxanames)) {
message("Ignoring argument taxanames since genetreedata supplied as quartet table.")
}
} else {
if ("multiPhylo" %in% class(genetreedata)) {
genetrees = genetreedata
} else {
if ("character" %in% class(genetreedata)) {
genetrees <- read.tree(genetreedata) #read gene trees
message(paste("Read", length(genetrees), "gene trees from file."))
} else {
stop("Data must be supplied as an object of class multiPhylo, character, or matrix.")
}
}
if (is.null(taxanames)) {
# if no taxa names specified,
taxanames = genetrees[[1]]$tip.label # ... get them from first tree
}
taxanames = sort(taxanames)
if (length(taxanames) <= 25) {
namelist = paste0(taxanames, collapse = ", ")
} else {
namelist = paste0(paste0(taxanames[1:25], collapse = ", "),
",...(see output table for full list)")
}
message("Analyzing ", length(taxanames), " taxa: ", namelist)
# tally quartets on gene trees
RQT = quartetTableResolved(quartetTable(genetrees, taxanames), omit = omit) # treat unresolved quartets as 1/3 of each resolution
}
Q = quartetTableDominant(RQT)
tree = QDS(Q,method=method)
if (metric==FALSE) {
tree$edge.length = NULL
} else {
tree = estimateEdgeLengths(tree, RQT, lambda = 0)
}
return(tree)
}
#############################################################
#' Compute Weighted Quartet Distance Consensus tree from gene tree data
#'
#' Compute the Weighted Quartet Distance Consensus \insertCite{YR19}{MSCquartets} estimate of a
#' species tree from gene tree data. This is a consistent estimator of the unrooted
#' species tree topology and all internal branch lengths.
#'
#' @details This function is a wrapper which performs the steps of reading in a collection
#' of gene trees, tallying quartets, estimating quartet internal branch lengths, computing the weighted
#' quartet distance between taxa, building
#' a tree, and adjusting edge lengths, to give a consistent estimate of the metric species tree in coalescent units
#' under the MSC.
#'
#' If the gene tree data indicates some quartets experienced little to no incomplete lineage
#' sorting, this algorithm tends to be less topologically accurate than \code{QDC}
#' (which infers no metric information) or \code{WQDCrecursive} (which gives better topologies,
#' and reasonably accurate lengths for short edges, though long edge lengths may still be unreliable).
#'
#' @references
#' \insertRef{YR19}{MSCquartets}
#'
#' @param genetreedata gene tree data that may be supplied in any of 3 forms:
#' \enumerate{
#' \item a character string giving the name of a file containing gene trees in Newick
#' \item a multiPhylo object containing gene trees
#' \item a resolved quartet table, as produced by \code{quartetTableResolved}
#' }
#' @param taxanames if \code{genetreedata} is a file or a multiPhylo object, a vector of a subset
#' of the taxa names on the gene trees
#' to be analyzed, if \code{NULL} all taxa on the first gene tree are used; if \code{genetreedata}
#' is a quartet table, this argument is ignored and all taxa in the table are used
#' @param method a distance-based tree building function, such as \code{fastme.bal} or \code{nj}
#' @param omit \code{TRUE} leaves out unresolved quartets, \code{FALSE} treats them as 1/3 of each resolution; ignored if
#' \code{genetreedata} is given as a resolved quartet table
#' @param terminal non-negative branch length to supply for terminal branches
#' whose length cannot be inferred by \code{WQDC}
#' @return an unrooted metric tree of type phylo
#' @seealso \code{\link{quartetTable}},
#' \code{\link{quartetTableResolved}},
#' \code{\link{quartetTableDominant}},
#' \code{\link{quartetWeightedDist}},
#' \code{\link{WQDCrecursive}},
#' \code{\link{WQDS}},
#' \code{\link{QDC}}
#'
#' @examples
#' gtrees=read.tree(file=system.file("extdata","dataGeneTreeSample",package="MSCquartets"))
#' tnames=taxonNames(gtrees)
#' stree=WQDC(gtrees,tnames[1:6])
#' write.tree(stree)
#' plot(stree)
#'
#' @export
WQDC = function(genetreedata,
taxanames = NULL,
method=fastme.bal,
omit = FALSE,
terminal = 1) {
if ("matrix" %in% class(genetreedata)) {
RQT = genetreedata
if (!is.null(taxanames)) {
message("Ignoring argument taxanames since genetreedata supplied as quartet table.")
}
} else {
if ("multiPhylo" %in% class(genetreedata)) {
genetrees = genetreedata
} else {
if ("character" %in% class(genetreedata)) {
genetrees <- read.tree(genetreedata) #read gene trees
message(paste("Read", length(genetrees), "gene trees from file."))
} else {
stop("Data must be supplied as an object of class multiPhylo, character, or matrix.")
}
}
if (is.null(taxanames)) {
# if no taxa names specified,
taxanames = genetrees[[1]]$tip.label # ... get them from first tree
}
taxanames = sort(taxanames)
if (length(taxanames) <= 25) {
namelist = paste0(taxanames, collapse = ", ")
} else {
namelist = paste0(paste0(taxanames[1:25], collapse = ", "),
",...(see output table for full list)")
}
message("Analyzing ", length(taxanames), " taxa: ", namelist)
# tally quartets on gene trees
RQT = quartetTableResolved(quartetTable(genetrees, taxanames), omit = omit) # treat unresolved quartets as 1/3 of each resolution
}
Q=quartetTableDominant(RQT, bigweights ="finite")
WQDC = WQDS(Q,method=method)
term_edges = which(WQDC$edge[, 2] <= length(taxanames)) #modify terminal edge lengths
WQDC$edge.length[term_edges] = terminal
return(WQDC)
}
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