## birthdeath.R (2009-05-10) ## Estimation of Speciation and Extinction Rates ## with Birth-Death Models ## birthdeath: standard model ## bd.ext: extended version ## Copyright 2002-2009 Emmanuel Paradis ## This file is part of the R-package `ape'. ## See the file ../COPYING for licensing issues. birthdeath <- function(phy) { if (!inherits(phy, "phylo")) stop('object "phy" is not of class "phylo"') N <- length(phy$tip.label) x <- c(NA, branching.times(phy)) dev <- function(a, r) { -2 * (lfactorial(N - 1) + (N - 2) * log(r) + r * sum(x[3:N]) + N * log(1 - a) - 2 * sum(log(exp(r * x[2:N]) - a))) } out <- nlm(function(p) dev(p[1], p[2]), c(0.1, 0.2), hessian = TRUE) if (out$estimate[1] < 0) { out <- nlm(function(p) dev(0, p), 0.2, hessian = TRUE) para <- c(0, out$estimate) se <- c(0, sqrt(diag(solve(out$hessian)))) } else { para <- out$estimate se <- sqrt(diag(solve(out$hessian))) } Dev <- out$minimum ## compute the 95 % profile likelihood CIs ## (not very clean... but seems to work -- EP 2003-03-29) CI <- matrix(NA, 2, 2) foo <- function(p) dev(p, para[2]) - 3.84 - Dev inc <- 1e-2 lo <- para[1] - inc up <- para[1] + inc while (foo(lo) < 0) lo <- lo - inc while (foo(up) < 0) up <- up + inc CI[1, 1] <- uniroot(foo, lower = lo, upper = para[1])$root if (CI[1, 1] < 0) CI[1, 1] <- 0 CI[1, 2] <- uniroot(foo, lower = para[1], upper = up)$root foo <- function(p) dev(para[1], p) - 3.84 - Dev lo <- para[2] - inc up <- para[2] + inc while (foo(lo) < 0) lo <- lo - inc while (foo(up) < 0) up <- up + inc CI[2, 1] <- uniroot(foo, lower = lo, upper = para[2])$root CI[2, 2] <- uniroot(foo, lower = para[2], upper = up)$root names(para) <- names(se) <- rownames(CI) <- c("d/b", "b-d") colnames(CI) <- c("lo", "up") obj <- list(tree = deparse(substitute(phy)), N = N, dev = Dev, para = para, se = se, CI = CI) class(obj) <- "birthdeath" obj } print.birthdeath <- function(x, ...) { cat("\nEstimation of Speciation and Extinction Rates\n") cat(" with Birth-Death Models\n\n") cat(" Phylogenetic tree:", x$tree, "\n") cat(" Number of tips:", x$N, "\n") cat(" Deviance:", x$dev, "\n") cat(" Log-likelihood:", -(x$dev)/2, "\n") cat(" Parameter estimates:\n") cat(" d / b =", x$para[1], " StdErr =", x$se[1], "\n") cat(" b - d =", x$para[2], " StdErr =", x$se[2], "\n") cat(" (b: speciation rate, d: extinction rate)\n") cat(" Profile likelihood 95% confidence intervals:\n") cat(" d / b: [", x$CI[1, 1], ", ", x$CI[1, 2], "]", "\n", sep = "") cat(" b - d: [", x$CI[2, 1], ", ", x$CI[2, 2], "]", "\n\n", sep = "") } bd.ext <- function(phy, S) { if (!inherits(phy, "phylo")) stop('object "phy" is not of class "phylo"') if (!is.null(names(S))) { if (all(names(S) %in% phy$tip.label)) S <- S[phy$tip.label] else warning('the names of argument "S" and the names of the tip labels did not match: the former were ignored in the analysis.') } N <- length(S) x <- branching.times(phy) x <- c(x[1], x) trm.br <- phy$edge.length[phy$edge[, 2] <= N] dev <- function(a, r) { -2 * (lfactorial(N - 1) + (N - 2) * log(r) + (3 * N) * log(1 - a) + 2 * r * sum(x[2:N]) - 2 * sum(log(exp(r * x[2:N]) - a)) + r * sum(trm.br) + sum((S - 1) * log(exp(r * trm.br) - 1)) - sum((S + 1) * log(exp(r * trm.br) - a))) } out <- nlm(function(p) dev(p[1], p[2]), c(0, 0.2), hessian = TRUE) if (out$estimate[1] < 0) { out <- nlm(function(p) dev(0, p), 0.2, hessian = TRUE) para <- c(0, out$estimate) se <- c(0, sqrt(diag(solve(out$hessian)))) } else { para <- out$estimate se <- sqrt(diag(solve(out$hessian))) } Dev <- out$minimum cat("\nExtended Version of the Birth-Death Models to\n") cat(" Estimate Speciation and Extinction Rates\n\n") cat(" Data: phylogenetic:", deparse(substitute(phy)), "\n") cat(" taxonomic:", deparse(substitute(S)), "\n") cat(" Number of tips:", N, "\n") cat(" Deviance:", Dev, "\n") cat(" Log-likelihood:", -Dev/2, "\n") cat(" Parameter estimates:\n") cat(" d / b =", para[1], " StdErr =", se[1], "\n") cat(" b - d =", para[2], " StdErr =", se[2], "\n") cat(" (b: speciation rate, d: extinction rate)\n") }