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Mark Wilkinson, of the Department of Zoology, The Natural History Museum, London, U.K. (marw  (at) nhm.ac.uk) has produced REDCON, version 3.0, a program to implement his method of reduced consensus trees. These find a tree with possibly fewer species that satisfies a strict or a majority rule consensus criterion. REDCON reads trees in PAUP* format. It is a DOS executable, and is available at his software Web site at http://www.nhm.ac.uk/research-curation/research/projects/software/mwphylogeny.html.


Mark Wilkinson, of the Department of Zoology, The Natural History Museum, London, U.K. (marw  (at) nhm.ac.uk) has produced TAXEQ3, a program to carry out Safe Taxonomic Reduction, which means dropping some species to get a set whose phylogenetic relationships are less ambiguous. The method is described in a paper: Wilkinson, M. 1995. Coping with abundant missing entries in phylogenetic inference using parsimony. Systematic Biology 44: 501-514. TAXEQ3 is distributed as a DOS executable with documentation and sample data set from his software Web site at http://www.nhm.ac.uk/research-curation/research/projects/software/mwphylogeny.html.


[RadCon icon] Joseph Thorley of Poisson Consulting, Nelson, B.C., Canada (info  (at) poissonconsulting.com) and Rod Page of the University of Glasgow have written RadCon, version 1.1.6, a program to compute consensus trees, supertrees, measures of the shape of trees, and to rearrange trees. It can compute strict, semi-strict, Adams, and majority-rule consensus trees, Reduced consensus trees, MRP supertrees, Cladistic Information Content, Leaf Stability, and Double Decay Analysis. It can also measure the shape and resolution of trees, prune and regraft leaves, add outgroups, and reroot trees. It is described in a paper: Thorley, J. L. and R. D. M. Page. 2000. RadCon: Phylogenetic comparison and consensus. Bioinformatics 16: 486-487. RadCon is a Mac OS executable for Mac OS 7.5 or later. RadCon is available at its web site at http://www.poissonconsulting.ca/index.php/software/ and its manual can also be downloaded or viewed from there.


Vladimir Makarenkov (makarenv  (at) magellan.umontreal.ca), then of the Département de Sciences Biologiques of the Université de Montréal, Québec and now of the Departement d'Informatique of the Université du Québec à Montréal and the Département de Sciences Biologiques of the Université de Montréal, Québec has written a program to compute the Robinson and Foulds topological distance. This is a distance measure between trees, counting the numbers of branches on the two trees that have no counterpart on the other tree. The algorithm used is described in a paper: Makarenkov, V., and Leclerc, B. 2000. An optimal way to compare additive trees using circular orders. Journal of Computational Biology 7: 731-744. The program is available as C source code and as DOS executable and a Mac OS executable from its web site at http://www.bio.umontreal.ca/Casgrain/en/labo/robinson_foulds.html.


[TopD/fMts icon]  Pere Puigbò Avalos of the Evolutionary Genomics Group, Biochemistry and Biotechnology Department of Rovira i Virgili University, Tarragona, Spain (ppuigbo (at) urv.cat) has produced TopD/fMts (TOPological Distance / From Multiple To Single), version 3.3d, program to calculate distances between trees. TopD calculates distances between trees by the following methods: split distance. nodal distance, disagreement, taxa in common, and quartets distance. fMtS is used with it to subsample gene families to get trees with one copy per species. TopD/fMts can also list the taxa that are responsible for the disgreement between two trees. It is described in the paper: Puigbò P., S. Garcia-Vallvé and J. O. McInerney. 2007. TOPD/FMTS: a new software to compare phylogenetic trees. Bioinformatics, published online on April 25, 2007. It is available as Perl script. It can be downloaded from its web site at http://genomes.urv.es/topd/


Chris Creevey, then of the Bioinformatics and Pharmacogenomics Laboratory at National University of Ireland Maynooth (and now of Teagasc (The Irish Agriculture and Food Development Authority) in Carlow, Ireland, has written Clann (the Irish word for "family") version 3.0.2, a program to computer supertrees. It implements most of the major supertree methods, including matrix representation by parsimony (MRP), methods involving distance matrices, quartets methods, and splits methods, and includes a bootstrap method that samples from among the input trees. Clann is described in a paper: Creevey, C. J. and J. O. McInerney. 2005. Clann: investigating phylogenetic information through supertree analyses. Bioinformatics 21: 390-392. It is available as Windows, Mac OS X, and Linux executables from its web site at http://bioinf.may.ie/software/clann/.


[SuperTree icon] Nicolas Salamin at the Department of Ecology and Evolution of the University of Lausanne, Switzerland has written SuperTree, version 0.85b, (not, of course to be confused with Supertree) a program to take a set of input trees and build a matrix representation of them that can be used to compute a supertree using the MRP (matrix representation by parsimony) method. The program does not do the parsimony reconstruction from this matrix itself. The options available are described in a paper: Salamin, N., T. R. Hodkinson, V. Savolainen. 2002. Building supertrees: An empirical assessment using the grass family (Poaceae). Systematic Biology 51: 136 - 150. SuperTree is a Java runtime executable which will work on Linux, Windows, and Mac OS X systems, as long as they have the Sun Java Runtime JRT system version 1.3 or greater. The Java executables are distributed in Windows and in Linux/Mac versions. The Java source code can be obtained from Salamin, if desired, by contacting him by email. The program and its distribution are briefly described at its site at http://www2.unil.ch/phylo/bioinformatics/supertree.html  The program can be obtained from Salamin by emailing him (wwwphylo (at) unil.ch).


Olaf Bininda-Emonds at the Institut für Biologie und Umweltwissenschaften of the Universität Oldenburg, Germany (olaf.bininda  (at)  uni-oldenburg.de) has produced a set of Perl scripts which, taken together, are a Supertree package which can make and examine supertrees using the program PAUP*. Although they are each downloadable separately, I will consider them here to be a single entity. They include, in rough order of the steps carried out:

Additional Perl scripts are also present to help with labeling the resultant supertrees and interconverting tree formats.

They are described in the papers:

As they are Perl scripts, they can be run on any system on which Perl has been made available. They can be downloaded (individually) from Bininda-Emonds's software web page at http://www.molekularesystematik.uni-oldenburg.de/33997.html


[Supertree icon]  Rod Page of the Environmental and Evolutionary Biology of the Institute of Biomedical and Life Sciences, University of Glasgow, U.K. (r.page (at) bio.gla.ac.uk) has written Supertree (not, of course to be confused with SuperTree) version 0.3, for constructing supertrees. Supertree is an experimental command line program for constructing supertrees. It implements Semple and Steel's algorithm MinCutSupertree (original and a modified version due to Page), as well as MRP coding. It can also compute cluster graphs. It is described in the paper: Page, R. D. M. 2002. Modified MinCut supertrees. pp. 537-551 in Algorithms in Bioinformatics: Proceedings of the Second International Workshop, WABI 2002, Rome, Italy, September 17-21, 2002. Springer-Verlag, Heidelberg. It is available as C source code, Mac OS X PowerMac executables and Mac OS 9 executables. To compile the source code, one needs a copy of the GTL Graph Template Library, which must be installed before compiling. Supertree can be downloaded from its web site at http://darwin.zoology.gla.ac.uk/%7Erpage/supertree/


Duhong Chen, then of the Computational Biology Laboratory of the Department of Computer Science of Iowa State University (duhong (at) iastate.edu) has written HeuristicMRF2 (Matrix Representation with Flipping version 2), a program to construct supertrees by the MRF method. Matrix representation with flipping (MRF) starts with a Matrix Representation by Parsimony matrix. The binary MRP matrix from a rooted input tree may be transformed into a subset of the columns of a rooted supertree by changing some of the 0's to 1's and 1's to 0's. Each change in the character state is a flip, and the minimum number of flips needed to transform the input tree into a supertree is the flip distance. The MRF heuristic seeks the rooted supertree(s) that minimizes the total flip distance from all input trees. It searches for rooted trees, not unrooted trees. It is described in the paper: Chen, D., L. Diao, O. Eulenstein, D. Fernández-Baca, and M. J. Sanderson. 2003. Flipping: A supertree construction method. Pp. 135-160 in Bioconsensus., ed. M. F. Janowitz, F.-J. Lapointe, F. R. McMorris, B. Mirkin, and F. S. Roberts. DIMACS series in discrete mathematics and theoretical computer science, Volume 61. American Mathematical Society, Providence, Rhode Island. It is available as C++ source code. It can be downloaded from its web site at http://genome.cs.iastate.edu/CBL/download/


Raul Piaggio of the Computational Biology Laboratory, Department of Computer Science at Iowa State University, Ames, Iowa (rpiaggio (at) iastate.edu) has written Quartet Suite version 1.0, a set of programs for computing supertrees and also distances between trees from quartets. Quartet Suite is a set of four programs that take input trees, break them down into the set of quartets implied by each of them, and construct a supertree based on these quartets. They can also compute a distance between trees from the sets of quartets they imply. The methods are described in the paper: Willson, S.J. 1999. Building phylogenetic trees from quartets by using local inconsistency measures. Molecular Biology and Evolution 16: 685-693. It is available as C++ source code, Windows executables and Powermac Mac OS X executables. It can be downloaded from its web site at http://genome.cs.iastate.edu/CBL/download/


Duhong Chen, Oliver Eulenstein, and David Fernández-Baca of the Computational Biology Laboratory of the Department of Computer Science at Iowa State University, Ames, Iowa (duhong (at) iastate.edu) have released Rainbow version beta 1.2, a toolbox for phylogenetic supertree construction and analysis. Rainbow provides a user-friendly environment in which scientists can utilize tools for building and analyzing supertrees. Rainbow provides a graphic user interface (GUI) to construct supertrees using several different methods. Currently these include matrix representation with flipping (MRF) matrix representation with parsimony (MRP), and the modified Mincut algorithm (MMC). Rainbow also provides tools to analyze the quality of the inferred supertrees. The methods are described in the paper: Chen, D., L. Diao, O. Eulenstein, D. Fernández-Baca, and M. J. Sanderson. 2003. Flipping: A supertree construction method. Pages 135-160 in Bioconsensus, ed. M. F. Janowitz, F.-J. Lapointe, F. R. McMorris, B. Mirkin, and F. S. Roberts. Volume 61 in DIMACS series in discrete mathematics and theoretical computer science, American Mathematical Society, Providence, Rhode Island. It is available as C++ source code, Windows executables, Linux executables and Powermac Mac OS X executables. It can be downloaded from its web site at http://genome.cs.iastate.edu/CBL/download/


Vincent Ranwez, Vincent Berry, Alexis Criscuolo, Pierre-Henri Fabre, Sylvain Guillemot, Celine Scornavacca and Emmanuel Douzery of the Institut des Sciences de L'evolution and the LIRMM at the Université Montpellier II, France (vberry (at) lirmm.fr) have produced PhySIC_IST (Phylogenetic Supertrees with Induction and non-Contradiction), a supertree program that uses an algorithm they developed. It carries out construction of supertrees using the non-contradiction property (PC) and the induction property (PI). The former requires that the supertree does not contain relationships that contradict one or a combination of the source topolo- gies, while the latter requires that all topological information contained in the supertree is present in a source tree or collectively induced by several source trees. The program also can collapse branches in the input trees that have bootstrap values that are smaller than a threshold set by the user. It is described in the papers:

It is available as Linux executables and Intel Mac OS X executables. It can be downloaded from its web site at http://www.atgc-montpellier.fr/physic_ist/binaries.php  It is available as a web server here. An earlier version PhySIC, is downloadable here at http://www.atgc-montpellier.fr/physic/binaries.php and it is also available as a web server here.


Ahmed Moustafa, then of the Computational Genetics program at the University of Iowa, Iowa City, Iowa (ahmed (at) users.sourceforge.net) and currently in Bioinformatics and Genomics at the Department of Biology and Biotechnology Graduate Program of the School of Science and Engineering, American University in Cairo, Egypt (ahmed (at) pobox.com) has written PhyloSort version 1.3, a Java tool for sorting phylogenies searching for user-specified subtrees that a particular monophyletic group. PhyloSort is for

It is available as Java source code and Java executables. It is described in the paper: Moustafa, A. and D. Bhattacharya. 2008. PhyloSort: A user-friendly phylogenetic sorting tool and its application to estimating the cyanobacterial contribution to the nuclear genome of Chlamydomonas. BMC Evolutionary Biology 8: 6. It can be downloaded from its web site at http://phylosort.sourceforge.net/


[CONSERVE icon here] The late (and lamented) Ross Crozier of the School of Tropical Biology, James Cook University, Townsville, Australia and Paul-Michael Agapow, then of the Department of Biology of Imperial College at Silwood Park, U.K. and more recently of the Health Protection Agency, U.K. (agapow  (at) agapow.net) have written CONSERVE, currently at version IV, 1.4b Beta, a program to use phylogenetic information to calculate biodiversity and test the feasability of conservation schemes. It measures the distinctiveness of species using genetic distances and also to test whether particular assemblages of populations preserve statistically significantly more biodiversity than other assemblages. Biodiversity is determined using GD (probability of more than one allele) or PD (length of evolutionary history) methods, from data in the form of unrooted trees produced in standard treefile format. It is available as executables for Mac OS X, for Windows, and for Linux from Agapow's web site at http://www.agapow.net/software/conserve. An earlier 680x0 Mac OS version (version III 3.2.2) is available here at http://www.bio.ic.ac.uk/evolve/software/conserve/index.html. (Here, here, and here are some of the many tributes to the late Ross Crozier).


Georg Weiller, of the Genomic Interactions Group at the Research School of Biological Sciences of the Australian National University, Canberra (georg.weiller  (at) rsbs.anu.edu.au) has released TreeDis version 2.0. TreeDis finds the patristic distances (total length of branches between all pairs of taxa in a phylogeny. It takes as input the tree file in Newick standard form or in the format for NJTREE. It is distributed as a DOS executable (a C++ source code version can also be obtained from Weiller). It is available from its web site at http://www.rsbs.anu.edu.au/Products&Services/BiotechnologyTransferUnit/tredis.asp.


Nicolas Bortolussi, Eric Durand, Michael Blum, and Olivier François, then of the Institut d'Informatique et Mathématiques Appliquées in Grenoble, France (Durand is now the the Department of Integrative Biology, University of California, Berkeley and his email address is eric.durand (at) berkeley.edu) have written apTreeshape (analyses of phylogenetic Treeshape), version 1.4-3, an R package for simulation and analysis of phylogenetic tree topologies using statistical indices . apTreeshape computes a variety of statistics and tests on tree shape. It is a companion library of the APE package. It provides additional functions for reading, plotting, manipulating phylogenetic trees. It also offers convenient web-access to public databases, and enables testing null models of macroevolution using corrected test statistics. Trees of class "phylo" (from the APE package) can be converted easily. It is available as an R package, Windows executables and Powermac Mac OS X executables. It can be downloaded from its web site at http://cran.r-project.org/web/packages/apTreeshape/index.html


[TreeStat icon]  Andrew Rambaut of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) and Alexei Drummond of the University of Auckland, New Zealand have written TreeStat (Tree Statistics), version 1.2, a phylogeny package with parsimony, distance and likelihood methods. TreeStat is an application that can process a set of trees in a PHYLIP or NEXUS format tree file and calculate a number of summary statistics for each. These are saved in a tab-delimited file for analysis in Tracer or statistics packages. A range of summary statistics are included:

It is available as Java executables, Windows executables and Mac OS X universal executables. It can be downloaded from
its web site at http://tree.bio.ed.ac.uk/software/treestat/


   Federico Plazzi, Ronald Robert Ferrucci, and Marco Passamonti of the Department of Experimental Evolutionary Biology of the University of Bologna, Italy (federico.plazzi (at) unibo.it) have released PhyRe (PHYlogenetic REpresentativeness), version 1.0, a package which aims to estimate taxonomic coverage of an ingroup. PhyRe implements a method to evaluate and assess taxon sampling in a phylogenetic study by means of Average Taxonomic Distinctness and related indices and algorithms. It was conceived as an a priori checkpoint to be performed before starting an evolutionary study, whether based on morphology or molecules. The method uses a tree of the group which is being investigated; PhyRe can also test the stability of results upon taxonomical revisions. It is described in the paper: Plazzi, F., R. R. Ferrucci and M. Passamonti. 2010. Phylogenetic Representativeness: A new method for evaluating taxon sampling in evolutionary studies. BMC Bioinformatics 11: 209. It is available as Python script and Windows executables. It can be downloaded from its web site at http://www.mozoolab.net/index.php/software-download.html


[CAIC icon here] Andrew Purvis of the Department of Biology, Imperial College, Silwood Park, U.K. (a.purvis (at) imperial.ac.uk) and Andrew Rambaut of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) have written CAIC (Comparative Analysis of Independent Contrasts), versions 2.6.9 and 3-Alpha. It carries out the contrasts method but with some modifications by others to cope with lack of resolution of the phylogeny. It is described in the paper: Purvis, A. and A. Rambaut. 1995. Comparative analysis by independent contrasts (CAIC): an Apple Macintosh application for analysing comparative data. Computer Applications in the Biosciences (CABIOS) 11: 247-251. It will run on Macintosh Mac OS, and is available free from CAIC's Web page http://www.bio.ic.ac.uk/evolve/software/caic/index.html. It is also available as an "alpha" release of an R package for CAIC that implements its methods is available at its page at R-Forge at http://r-forge.r-project.org/projects/caic/


Emília Martins (emartins  (at) indiana.edu), of the Department of Biology of the University of Indiana, Bloomington, Indiana, has released version 4.6b of COMPARE, a package of programs for comparative methods analysis. COMPARE includes various programs for conducting statistical analyses of comparative data in a phylogenetic context. At the moment, it includes programs to compute independent contrasts, do spatial autocorrelation analyses, sum of squares parsimony, generate random data, trees and/or branch lengths, and various other things. COMPARE is written in Java and is available both as standalone Java (including source code) and as a Java application that is provided by their site to your browser. It requires a Java runtime environment. COMPARE is available from its web site at http://compare.bio.indiana.edu/.


Theodore Garland, Jr., of the Department of Biology of the University of California, Riverside (tgarland  (at) ucr.edu) and his colleagues (Jason A. Jones, Allan W. Dickermann, Peter E. Midford, and Ramon Diaz-Uriarte) have developed PDAP version 6.0, Phenotypic Diversity Analysis Programs, a series of DOS programs to perform various comparative analyses. At present, the following phylogenetically based statistical methods are included: independent contrasts, squared-change parsimony reconstructions of ancestral states and estimation of evolutionary correlations, and phylogenetic analysis of covariance via computer-simulated (Monte Carlo) null distributions. PDTREE can also read, write, and edit trees. PDAP distribution is described in a web page at http://www.biology.ucr.edu/people/faculty/Garland/PDAP.html. The original published description of PDAP is the paper: Garland, T., Jr., A. W. Dickerman, C. M. Janis, and J. A. Jones. 1993. Phylogenetic analysis of covariance by computer simulation. Systematic Biology 42:265-292. The methods used are described in a number of recent papers by these authors, including:

PDAP is distributed by email of a self-extracting executable file, obtainable for free (contact Garland by e-mail at the above address). Alternatively, a DOS disk can be mailed.


Liam Revell, then of the Department of Organismic and Evolutionary Biology of Harvard University (currently at the National Evolutionary Synthesis Center, Duke University, Durham, North Carolina, email address (lrevell (at) nescent.org) has released IDC (IndepenDent Contrasts program), version 1, a program for the calculation of phylogenetically indepedent contrasts. This program calculates contrasts for multiple trees, multiple datasets, or both. It also returns a VCV matrix and the correlation matrix of the independent contrasts. It is available as C source code and Windows executables. It can be downloaded from its web site at http://anolis.oeb.harvard.edu/~liam/programs/


[PI icon] Ehab Abouheif (ehab.abouheif  (at) staff.mcgill.ca) of the Department of Biology, McGill University, Montréal, Québec has written (together with J. Reeve) Phylogenetic Independence version 2.0. It carries out Abouheif's Test For Serial Independence (TFSI) on continuously valued characters and his Runs Test on discretely valued characters. These are described in his paper: Abouheif, E. 1999. A method to test the assumption of phylogenetic independence in comparative data. Evolutionary Ecology Research 1: 895-909. The methods employed are validate in the paper: Pavoine, S., S. Ollierb, D. Pontiera, and D. Chessela. 2008. Testing for phylogenetic signal in phenotypic traits: New matrices of phylogenetic proximities. Theoretical Population Biology 73: 79-91. The program is available as Windows and Linux executables at its web site at http://biology.mcgill.ca/faculty/abouheif/.


Mark Pagel and Andrew Meade of the School of Biological Sciences, of the University of Reading, Reading, U.K. (m.pagel (at) reading.ac.uk) have released BayesTraits, version 1.0, a Bayesian package of programs to analyze state evolution of discrete and continuous traits. It is performs analyses of trait evolution among groups of species for which a phylogeny or sample of phylogenies is available. It incoporates their earlier and separate programs Multistate, Discrete and Continuous. BayesTraits can be applied to the analysis of traits that adopt a finite number of discrete states, or to the analysis of continuously varying traits. Hypotheses can be tested about models of evolution, about ancestral states and about correlations among pairs of traits. Parts of the package are described in these papers:

BayesTraits is available as Windows executables, Linux executables, Powermac Mac OS X executables, and Intel Mac OS X executables. It can be downloaded from its web site at http://www.evolution.rdg.ac.uk/BayesTraits.html


Daniel Barker and Mark Pagel, respectively of the School of Biology of the University of St. Andrews and the School of Biological Sciences of the University of Reading, U.K. (db60 (at) st-andrews.ac.uk) have written bms_runner version 1.3, a Perl script to use BayesTraits to detect correlations in gain and loss of genes. It uses the separately distributed program BayesTraits to predict functional links among genes, on the basis of correlated gain and loss of genes from species' genomes. bms_runner helps configure BayesTraits on the basis of user-supplied data, and launches BayesTraits repeatedly to seek correlated gain and loss within pairs of genes. The methods are described in the paper: Barker, D., Meade, A. and Pagel, M. 2007. Constrained models of evolution lead to improved prediction of functional linkage from correlated gain and loss of genes. Bioinformatics 23: 14-20. It is available as Perl script. Use of bms_runner requires that BayesTraits also be installed. It can be downloaded from its web site at http://eggg.st-andrews.ac.uk/bms_runner


Aaron King and Marguerite Butler of the Department of Ecology and Evolutionary Biology of the University of Michigan, Ann Arbor, Michigan and the Department of Zoology of the Hawaii at Manoa (aaron.king (at) umich.edu and mbutler (at) hawaii.edu) have written OUCH (Ornstein-Uhlenbeck models models for phylogenetic Comparative Hypotheses), version 2.7-1, an R package using the Ornstein-Uhlenbeck model for comparative methods tests. The package fits different versions of Ornstein-Uhlenbeck models to comparative data, as described by Thomas Hansen. It is described in the paper: Butler, M. A. and A. A. King, 2004. Phylogenetic comparative analysis: a modeling approach for adaptive evolution. American Naturalist 164: 683-695. It is available as an R package, and requires the R statistical computing enironment to be available. It can be downloaded from its web site at http://tsuga.biology.lsa.umich.edu/king/ouch/ and is also available at its web site at the CRAN-R archive site for R packages at http://cran.at.r-project.org/web/packages/ouch/


Chunghau Lee and Todd Oakley of the Ecology, Evolution and Marine Biology Department of the University of California at Santa Barbara, Santa Barbara, California (chunghaulee (at) gmail.com) and (oakley (at) lifesci.ucsb.edu) have released CoMET (COntinuous-character Model Evaluation and Testing), version of 2007-09-22, a Mesquite module for computing likelihoods for a given tree with Brownian motion models. COMet is a module for the Mesquite Project. Give it a tree topology as a starting point and some phenotypic data, and CoMET tells you the likelihoods of the data evolving through nine different evolutionary models, including both gradualist and punctuational models. It is described in the paper: Lee, C., S. Blay, A. Ø. Mooers, A. Singh, and T. H. Oakley. 2006. CoMET: A Mesquite package for comparing models of continuous character evolution on phylogenies. Evolutionary Bioinformatics Online 2: 193-196. CoMET requires the Mesquite Java framework for phylogenies. It is available as Java source code and Java executables. It can be downloaded from its web site at http://www.lifesci.ucsb.edu/eemb/labs/oakley/software/comet.htm


Brian O'Meara of the Center for Population Biology of the University of University of California, Davis. (bcomeara (at) ucdavis.edu) has released Brownie version 2.1.2, a program for analyzing rates of continuous character evolution. Brownie looks for substantial rate differences in different parts of a tree using likelihood ratio tests and Akaike Information Criterion (AIC) statistics. Brownie 2.1.2 can read Nexus tree and data files and can perform analyses across a set of weighted input trees (such as trees weighted by posterior probabilities or bootstrap values) in order to deal with tree uncertainty. Brownie deals with subtrees (using the censored test of O'Meara et al) and asks whether the subtrees differ in rates of evolution. An earlier version, Brownie 1.0, is written as a series of MATLAB routines and can be used on systems that have MATLAB installed. It is described in the paper: O'Meara, B.C., C. Ané, M.J. Sanderson, and P.C. Wainwright. 2006. Testing for different rates of continuous trait evolution using likelihood. Evolution 60(5): 922-933. It is available as C++ source code, Windows executables and Mac OS X universal executables. It can be downloaded from its web site at http://www.brianomeara.info/brownie/


Emmanuel Paradis (Emmanuel.Paradis (at) ird.fr) of the Institut de Recherche pour le Développement, Paris, France, has written APE, version 2.6-1, (Analysis of Phylogenetics and Evolution) a package in the R statistical and graphical language which carries out a variety of phylogeny analyses, including computation of distances from sequences and gene frequencies, comparative methods, analyses of diversification, computation of minimum spanning trees, and estimation of rates of evolution and smoothing of rates in neighboring lineages. Some code was contributed by a number of other people, including Korbinian Strimmer, Julien Claude, Gangolf Jobb, Rainer Opgen-Rhein, Julien Dutheil, Yvonnick Noel, and Ben Bolker. APE is described in a paper: Paradis, E., J. Claude and K. Strimmer. 2004. APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20: 289-290. APE can be downloaded from its page at the CRAN-R archive site of R programs at http://cran.r-project.org/web/packages/ape/index.html; it is described and some additional documentation links given at its web site at http://ape.mpl.ird.fr/


Liang Liu and Lili Yu of the Department of Agriculture and Natural Resources of the Delaware State University (lliu (at) desu.edu) has released Phybase (An R package for species tree analysis), version 1.1, a phylogeny package for manipulating, estimating, and summarizing phylogenetic trees, especially species trees. This package provides functions to read, write, manipulate, estimate, and summarize phylogenetic trees including species trees which contains not only the topology and branch lengths but also population sizes. The input/output functions can read tree files in which trees are presented in parenthetic format. The trees are read in as a string and then transformed to a matrix which describes the relationship of nodes and branch lengths. The nodes matrix provides an easy access for developers to further manipulate the tree, while the tree string provides interface with other phylogenetic R packages such as "ape". The input/output functions can also be used to change the format of tree files between NEXUS and PHYLIP. Some basic functions have already been established in the package for manipulating trees such as deleting and swapping nodes, rooting and unrooting trees, changing the root of the tree. The package also includes functions such as "consensus", "coaltime, "popsize", "treedist" for summarizing phylogenetic trees, calculating the coalescence time, population size, and tree distance. It is described in the paper: Liu, L., and L. Yu. 2010. Phybase: an R package for species tree analysis. Bioinformatics 26: 962-963. It is available as an R package. Instructions for downloading it are available at its web site at http://cars.desu.edu/faculty/lliu/research/phybase.html


Ramón Díaz-Uriarte, of the Bioinformatics Unit of the Spanish National Cancer Center (CNIO), Madrid, Spain (rdiaz (at) ligarto.org) and Theodore Garland, Jr, of the Department of Biology of the University of California, Riverside (garland  (at) ucr.edu have released PHYLOGR, version 1.0.7. This is a package of programs in the R statistical language (and also available as a Windows executable) to carry out comparative methods analyses, particularly ones using the Generalized Least Squares method. The package is distributed through the web site of the Cran-R (Comprehensive R Archive Network) project at http://cran.r-project.org/web/packages/PHYLOGR/index.html


Jason Pienaar, Thomas F. Hansen, and Steven Hecht Orzack of (respectively) the Department of Molecular Genetics and Microbiology of the University of Florida, Gainesville, Florida (jasonpienaar (at) ufl.edu), the Center for Ecological and Evolutionary Synthesis at the University of Oslo, Norway (t.f.hansen (at) bio.uio.no) who is also of the Department of Biological Sciences at Florida State University, Tallahassee, Florida (thomas.hansen (at) bio.fsu.edu), and the Fresh Pond Research Institute, 173 Harvey Street, Cambridge, Massachusetts 02140 (orzack (at) freshpond.org) have written SLOUCH (Stochastic Linear Ornstein-Uhlenbeck models for Comparative Hypotheses), version 1.0, an R package for comparative methods analysis using an Ornstein-Uhlenbeck model. SLOUCH allows the user to estimate 1) the evolutionary and optimal regressions between a predictor and a response trait, and 2) phylogenetic inertia. These quantities are estimated jointly by a comparative method based upon an Ornstein-Uhlenbeck model of adaptive evolution in which a single trait adapts to an optimum that is influenced by one or more continuous, randomly changing predictor variables. The methods are described in the paper: Hansen, T. F., J. Pienaar, and S. H. Orzack. 2008. A comparative method for studying adaptation to a randomly evolving environment. Evolution 62: 1965-1977. It is available as an R package. It can be downloaded from its web site at http://freshpond.org/software/SLOUCH/


Klaus Schliep, then at Massey University, Palmerston North, New Zealand (K.P.Schliep (at) massey.ac.nz) has written phangorn version 1.1-2, an R package to do phylogenetic analysis using parsimony, distance, likelihood and Hadamard methods. phangorn can carry out parsimony methods, the neighbor-joining distance method, maximum likelihood, and both Hadamard and distance Hadamard methods. It can also compute tree distances. It is available as an R package. phangorn requires that the APE and stats packages also be installed. It can be downloaded from its web site at http://cran.r-project.org/web/packages/phangorn/index.html


Ted Garland of the Department of Biology of the University of California at Riverside (tgarland (at) citrus.ucr.edu) distributes PHYSIG (PHYlogenetic SIGnal), a MATLAB package of modules for comparative methods analysis. PHYSIG is a package of modules in MATLAB that includes tests for phylogenetic signal in continuous-character data, tests for the attraction to the mean in an Ornstein-Uhlenbeck model, and tests of evolutionary covariation between characters by a Generalized Least Squares method. It can be obtained by emailing Garland at the above address (it is not available by web download). The methods are described in the paper: 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. It is available as a MATLAB package, which is described at its web site at http://biology.ucr.edu/people/faculty/Garland/PHYSIG.html


[Cactus-Pie icon]   Dylan Schwilk of the Department of Biology, Texas Tech University, Lubbock, Texas (dylan.schwilk (at) ttu.edu) has written Cactus-Pie (Python version of CACTUS: Comparative Analysis of Continuous Traits Using Statistics), version 0.3.1, a program for comparative analysis of continuous traits. Cactus-pie is a version of CACTUS written in Python. It consists of a front end program and a number of back-end modules. They include programs that calculate independent contrasts, provide the Divergence Order Test (DOT) and the Synchronized Changes Test (SvS), assign branch lengths to a phylogeny according to several different possible algorithms, and label clades in a phylogeny as defined by a taxa list in an auxilary file. It is available as a Python script which can run on computers that have the Python language, which is widely available on Linux and Unix systems and is standard on Mac OS X. Cactus-Pie can be downloaded from its web site at http://www.pricklysoft.org/software/cactus-pie.html  An earlier version which is a Windows executable (using graphical libraries that may not work on Windows XP or larer) is also available there at http://www.pricklysoft.org/software/cactus.html


Liam Revell of the Department of Biology, University of Massachusetts, Boston (lrevell (at) nescent.org) has released pcca (Phylogenetic Canonical Correlation Analysis), Beta version, a program for phylogenetic canonical correlation analysis. pcca uses a phylogenetic tree and measurements for an arbitrary number of continuous characters to perform a PGLS transformation of the data and then calculate canonical scores, weights, and correlations, and conduct hypothesis tests about the canonical correlations. Canonical correlation analysis (CCA) is a procedure using two sets of variables and calculating the linear combinations of each that have highest correlation with each other. The program can also estimate a multivariate version of Pagel's lambda transformation of time, and perform the analyses under that transformation. It is described in the paper: Revell, L. J. and A. S. Harrison. 2008. PCCA: A program for phylogenetic canonical correlation analysis. Bioinformatics 24: 1018-1020. pcca is available as Windows executables, Linux executables and Mac OS X universal executables. It can be downloaded from its web site at http://anolis.oeb.harvard.edu/~liam/programs/


[Treescan icon] David Posada (dposada  (at) uvigo.es), of the Universidad Vigo, Spain, and Taylor Maxwell and Alan Templeton, of the Department of Biology of Washington University, Saint Louis, Missouri (temple_a  (at) biology.wustl.edu) have written TreeScan, version 1.0. This is a program to test the association of continuous quantitative characters with a tree of haplottypes. It is described in a paper: Templeton, A. R., T. Maxwell, D. Posada, J. H. Stengåd, . Boerwinkle, and C. F. Sing. 2005. Tree scanning: a method for using haplotype trees in phenotype/genotype association studies. Genetics 169: 441-453. Treescan is provided in C source code, and also as a DOS and Windows executable or a Mac OS X executable. It is available from its web site at http://darwin.uvigo.es/software/treescan.html


Dolph Schluter (schluter  (at) zoology.ubc.ca) of the Department of Zoology and Biodiversity Research Centre University of British Columbia, in Vancouver, Canada, has released ANCML, a program which estimates ancestor states for a continuous trait, and provides a "standard error" for the marginal distribution of each estimate. The method is described in Schluter, D., T. Price, A. Ø. Mooers and D. Ludwig. 1998. Likelihood of ancestor states in adaptive radiation. Evolution 51: 1699-1711. The method assumes a Brownian motion model for the evolution of the trait. ANCML was written by modifying the program CONTRAST in PHYLIP version 3.5, and it uses similar input conventions. ANCML is available from its web page at http://www.zoology.ubc.ca/~schluter/software.html. It is available as generic C source code, as a DOS executable, as a Mac OS X executable, and as a SunOS executable.


Bill Bruno (billb  (at) lanl.gov) of the Theoretical Biology and Biophysics Group T10, Los Alamos Scientific Laboratory, has produced RIND, (Reconstructed INDependence), a program which takes a tree supplied by the user, or uses a distance method of the users choosing (one which can be found in PHYLIP), and computes a maximum likelihood estimate of the number of times each residue in aligned protein sequences was replaced in each position. The method is described in: Bruno, W. J. 1996, Modeling residue usage in aligned protein sequences via maximum likelihood Molecular Biology and Evolution 13: 1368-1374. RIND is available from its web site at http://www.t10.lanl.gov/billb/rind/ as C source code for a Unix environment, and assumes that PHYLIP is also installed.


Steve Woolley, then of the Department of Computer Science, Brigham Young University, Provo, Utah, and now at the Department of Biology, Washington University, St. Louis, Missouri (stevenwoolley (at)  wustl.edu), together with Justin Johnson, Matthew Smith, Keith Crandall, and David McClellan of the Department of Integrative Biology, Brigham Young University (whose email address is David_McClellan  (at)  byu.edu) have released TreeSAAP version 3.2, (Selection on Amino Acid Properties), a program that analyzes the frequency of change in various properties of amino acids in sequences evolving on well-corroborated phylogenies supplied by the user. TreeSAAP is given nucelotide sequences and a user-defined tree, and optimizes the placement of changes on the tree. It then calculates measure of departure of these changes from models of uniform neutral substitution, with respect to different properties of the amino acids. The methods are described in a paper: Woolley, S., J. Johnson, M. J. Smith, K. A. Crandall, and D. A. McClellan. 2003. TreeSAAP: Selection on Amino Acid Properties using phylogenetic trees. Bioinformatics 19: 671-672. The program is a Java application. It is distributed as Windows executables, Mac OS X executables, and as Java source code from its web site at http://dna.cs.byu.edu/treesaap/


  Marcin Joachimiak (marcin  (at) compbio.berkeley.edu) of the Computational Genomics Research Group in the Department of Plant and Microbial Biology, University of California, Berkeley, has released Jevtrace version 3.12b. This is a Java package to analyze the distribution of amino acid changes across a phylogeny, and use protein structures to identify based on their distribution in the tree and in the protein structure. The program reads a multiple sequence alignment, a tree, and can also read a PDB protein structure. It displays the tree on either a branch length scale or a sequence similarity scale and allows the user to select clades of interest. The residues conserved in these clades and differing between them are then found and can then be viewed on the three-dimensional structure of the protein. Jevtrace is described in electronic publication: Joachimiak, M. P. and F. E. Cohen. 2003. JEvTrace: refinement and variations of the evolutionary trace in JAVA. Genome Biology 3 (12) http://genomebiology.com/2002/3/12/research/0077. The Jevtrace web page is available at http://compbio.berkeley.edu/people/marcin/jevtrace/. The software cannot be directly downloaded from there: the user should fill out the Academic Software License form and submit it. Distribution is free for academic institutions (prices for others are not stated). The user will be emailed a link to download the software.


Xun Gu, of the Department of Genetics, Development and Cell Biology and the Center for Bioinformatics and Biological Statistics at Iowa State University, Ames, Iowa (xgu  (at) iastate.edu) and Kent Vander Velden, then at that University, have released DIVERGE version 2.0. DIVERGE reads protein sequences and either infers a tree by Neighbor-Joining or lets you read in a tree that you supply. It then allows you to define two clades in the tree, and tests whether the pattern of rates of change at different sites differ in these two clades. The statistical method used, which is a likelihood ratio test based in a probabilistic model, is given in the paper: Gu, X. 1999. Statistical methods for testing functional divergence after gene duplication. Molecular Biology and Evolution 16: 1664-1674. The program is downloadable as Windows executables from the Gu laboratory software web site at http://xungulab.com/software.html. An earlier version 1.0.4, which has Windows and Linux executables, is also available there.


Mark Wilkinson, of the Department of Zoology, The Natural History Museum, London, U.K. (marw  (at) nhm.ac.uk) has written PICA, version 4.0, a package of programs for character weighting and randomization tests for compatibility analysis for 0/1 or multistate characters. These carry out a variety of tests for nonrandomly compatible characters and include methods developed by Sharkey, Le Quesne, Meacham and Alroy. They include ability of process data that reflect the splits method of Bandelt and Dress. The programs are available as a package of DOS executables, from his software web site at http://www.nhm.ac.uk/research-curation/projects/software/mwphylogeny.html .


Christopher Meacham, of the Berkeley Phylogenomics Group, and also affiliated with the University Herbarium, University of California, Berkeley has produced COMPROB, a Pascal program to compute probabilities that characters would be compatible at random, thus telling us which clique is "most surprising". He can be contacted as meacham  (at) berkeley.edu about receiving a copy. The program is free.


John Armstrong, Adrian Gibbs, R. Peakall and George Weiller, (John.Armstrong  (at) anu.edu.au) of Mark Gibbs's group at the School of Botany and Zoology of the Australian National University, Canberra, have produced RAPDistance version 1.04, a package for DOS for computing distance matrices for RAPD analyses. Version 2.00 was produced but seems not be currently available. Version 1.04 has slightly more functionality but cannot handle data sets as large as version 2.00. RAPDistance has a comprehensive range of options for creating data files, editing them and using application programs to analyse them. It can export data sets in format of several other packages. RAPDistance 1.04 is available free at a web page at http://www.rsbs.anu.edu.au/Products&Services/BiotechnologyTransferUnit/rapdistance.php


Peter Reeves, L. Farnell, and Ruiting Lan, then of the School of Molecular and Microbial Biosciences at Sydney University, Australia, have produced MULTICOMP, a program for computing various distances from sequence data. It can also do sequence format conversions, compute various descriptive statistics on the sequences, and can submit the sequences to two programs from PHYLIP. It is described in a paper: Reeves, P. R., L. Farnell and R. Lan. 1994. MULTICOMP: a program for preparing sequence data for phylogenetic analysis. Computer Applications in the Biosciences (CABIOS) 10: 281-284. It is available as SunOS executables and as Linux executables at a web site at ttp://www.emi.unsw.edu.au/~lanlab/public/multicomp/download.htm in the lab of Ruiting Lan at the University of New South Wales, Australia. Lan may be emailed to obtain the C source code, at (rlan (at) unsw.edu.au).


  Stuart Ray of the Division of Infectious Diseases at the Johns Hopkins University School of Medicine, Baltimore, Maryland (sray (at) jhmi.edu) have produced NimbleTree version 2.6, a program that submits data sets to PAUP* or PHYLIP. NimbleTree reads a variety of different sequence alignment formats, and allows you to more easily submit the resulting data sets to PHYLIP or to PAUP*. For PAUP* you need to have that program already installed on your computer. Some PHYLIP source code from its version 3.5p is included in NimbleTree (with my agreement). (I am not quite sure which methods from PHYLIP are available in this program). It is available as Windows executables. It can be downloaded from http://sray.med.som.jhmi.edu/SCRoftware/nimbletree/


Microsat versiobn 1.5, by Eric Minch, then of the Department of Human Genetics, Stanford University, Stanford, California (his email address is now eric.minch  (at) lionbioscience.com) is a program for calculating distances from microsatellite data. It uses the methods developed by David Goldstein et. al., and presented in their papers of 1995 in Proceedings of the Natonal Academy of Sciences USA 92: 6720-6727 and Genetics 139: 463-471. The distance is based on the mean microsatellite array size, implementing the "Δμ" distance that they defined, which corrects for within-population variability and provides a distance that is independent of population size. It also calculates a number of other microsatellite-based genetic distances. It is available for free from a page in Luca Cavalli-Sforza's lab web site at http://hpgl.stanford.edu/projects/microsat/. The program is written in ANSI C. Source code is available there, and so are executables for DOS and Mac OS.


Daniel Dieringer (daniel.dieringer  (at) boku.ac.at) and Christian Schlötterer of the University of Vienna have produced MSA (MicroSatellite Analyzer) version 4.05, a program for handling large microsatellite data sets. It can calculate many descriptive statistics for these data sets, can convert the data into a variety of file formats for other programs, and can also calculate a variety of distance measures. It is described in a paper: Dieringer, D., and C. Schlötterer. 2003. Microsatellite analyser (MSA): a platform independent analysis tool for large microsatellite data sets. Molecular Ecology Notes 3: 167-169. MSA is available as source code in C plus executables for Linux, Windows (and DOS), Mac OS, and Mac OS X from its web site at http://i122server.vu-wien.ac.at/MSA/MSA_download.html.


Georg Weiller, of the Genomic Interactions Group at the Research School of Biological Sciences, Australian National University, Canberra, Australia (georg.weiller  (at) anu.edu.au) has produced DIPLOMO (DIstance PLOt MOnitor) version 1.03. It compares different distance measures with each other by displaying them as a scatter plot. It then helps one instantly identify all individual comparisons within the plot. individual taxa can be excluded or included in the plots, DIPLOMO enables you to see whether different taxa have different mutational characteristics (such as more having relatively more transitions in some lineages), and whether different distance measures correlate. The program takes as input a file with several different distance matrices. This file is in a simple format which can readily be produced by editing distance matrices produced by other packages. It is described in a publication: Weiller, G. F. and A. Gibbs. 1995. DIPLOMO: The tool for a new type of evolutionary analysis. CABIOS 11: 535-40. It runs on DOS or in a Command Tool on Windows. It can be obtained from its web site at http://www.rsbs.anu.edu.au/Products&Services/BiotechnologyTransferUnit/diplomo.php.


  Oclair Prado (oclair  (at) cpqd.com.br) and Fernando Van Zuben (vonzuben  (at)  dca.fee.unicamp.br) of the Department of Computer Science of the University of Campinas, Brazil released the Phylogenetic Tree Project (PTP) genetic algorithms toolbox, version 1.0. It infers phylogenies by maximum likelihood or a distance matrix method, using an evolutionary computation strategy which represents a phylogeny by a genotype, simulates natural selection to choose among phylogenies and recombination to rearrange phylogenies. It is a Windows executable, which can be downloaded by ftp from file PTP.zip at ftp.dca.fee.unicamp.br in folder /pub/docs/vonzuben/oclair.


MetaPIGA icon   Alan R. Lemmon, of the Department of Scientific Computing and the Department of Biological Sciences at Florida State University, Tallahassee, Florida (alemmon  (at) evotutor.org) and Michel Milinkovich (mcmilink  (at) ulb.ac.be) of the Unit of Evolutionary Genetics, Institute of Molecular Biology and Medicine, Free University of Brussels, Belgium have written MetaPIGA 2 (Phylogeny Inference using the MetaGA), a program searching for maximum likelihood phylogenies using a genetic algorithm with metapopulations. This allows different populations in the genetic algorithm to arrive at different solutions, which can then be combined by migration and recombination. It is said to make possibe effective maximum likelihood inference of phylogenies for data sets with hundreds of species. It allows rate variation among sites, different substitution models, partitioning of data, and LRT, AIC amd BIC tests. MetaPIGA is described in two papers:

It is available as Java code from its web site at http://www.metapiga.org. The earlier version, MetaPIGA 1, is also available there.


  Derrick Zwickl of the Department of Ecology and Evolution of the University of Kansas, Lawrence, Kansas (zwickl (at) ku.edu) released GARLI (Genetic Algorithm for Rapid Likelihood Inference), version 1.0, a program using a genetic algorithm to search for maximum likelihood phylogenies. GARLI uses a genetic algorithm to perform heuristic phylogenetic searches under the General Time Reversible (GTR) model of nucleotide substitution and its submodels, with or without gamma distributed rate heterogeneity and a proportion of invariant sites. It can also analyze protein sequences using amino acid models and codon-based models. It can read NEXUS and PHYLIP format sequence files. An MPI algorithm is included for use on parallel computing clusters (the parallel version seeks to perform a more thorough tree search and does not reduce runtimes. The program was written when Zwickl was a graduate student at the Department of Zoology of the University of Texas at Austin. It is described in his thesis: Zwickl, D. J. 2006. Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. dissertation, The University of Texas at Austin. It is available as C++ source code, Windows executables and Mac OS X universal executables. It can be downloaded from its Google Code web page at http://code.google.com/p/garli/. An older version, called GARLI-PART version 0.97, which can handle partitioned models, is also available at its Wiki site at the National Evolutionary Synthesis Center at https://www.nescent.org/wg_garli/Partition_testing_version An even older version of GARLI, 0.951, is available here at http://www.bio.utexas.edu/faculty/antisense/garli/Garli.html.


MUST, version 5.15, a package of sequence management programs, was developed by Hervé Philippe (herve.philippe (at) umontreal.ca), of the Departement de Biochimie Université de Montréal, Québec. It intended as complementary to existing phylogeny and alignment programs and can produce output files in the formats of PHYLIP, PAUP*, Hennig86, and CLUSTAL. It contains a variety of sequence input, editing, checking, and storage functions, as well as a sequence editor and a phylogeny plotter. It also allows further analyses of the results from these phylogeny programs. The original version of MUST is described in a paper: Philippe, H. 1993. MUST, a computer package of management utilities for sequences and trees. Nucleic Acids Research 21: 5264-5272. MUST is available as a Linux/Unix program (including source code that can be compiled on Mac OS X as well) or an older and not recently-maintained DOS program (which can also be run in a command tool window on Windows) from its web page at http://megasun.bch.umontreal.ca/Software/HPLab/must/must.html


Steve Smith, formerly of the Harvard Genome Laboratory, wrote an X-Windows interactive sequence editor, GDE (Genetic Data Environment), version 2.2, which allows the user to edit sequences and align them by hand, and to select subsets of sites and sequences and call a variety of analysis proprams including ClustalV and many of the PHYLIP 3.5 programs. The GDE 2.2 system will run on Unix or Linux systems using the X windowing system. It also includes the TreeTool tree-plotting program (see below). GDE has been described in two papers:

GDE is free and is available as source code in C and as executables at these places: GDE is the precursor of the SeqLab interface used with the (now unavailable) GCG Wisconsin Package of sequence search and analysis programs.


Brian Fristensky of the Department of Plant Science of the University of Manitoba, Winnipeg, Manitoba, Canada (frist (at) cc.umanitoba.ca) distributes BIRCH (BIological Research Computer Hierarchy), version 2.1, a sequence management and submission system with molecular databases. BIRCH consists of scripts and programs for a wide variety of analyses. It uses the GDE system for sequence editing and submission to programs. Some of the phylogeny programs (about 13 programs or packages for phylogenies) include many from PHYLIP, fastDNAml, MrBayes, Phylo_win, ATV, and ClustalW. BIRCH when installed also includes copies of molecular databases; the user is instructed on how to download these from their distribution sites. BIRCH is written in C and in Python and Java. It is distributed in source code and can be compiled on Linux and Solaris. The programs are distributed as binaries. The main BIRCH web site is at http://home.cc.umanitoba.ca/~psgendb/. It is described in

BIRCH can be downloaded from its download web site at http://home.cc.umanitoba.ca/%7Epsgendb/downloads.html


Jongsik Chun of the School of Biological Sciences, Seoul National University, Korea (jchun  (at) snu.ac.kr) has released PHYDIT version 3.1. PHYDIT is a sequence alignment environment that can align either nucleotide or protein sequences manually or using pairwise methods, and contains some PHYLIP programs to do either neighbor-joining or least-squares phylogenies on the alignments. PHYDIT is available as a Windows executable. It is available free for noncommercial use. The user needs to register to download. The PHYDIT web page is http://plaza.snu.ac.kr/~jchun/phydit/.


[SeqPup icon] [SeqPup Java icon] Don Gilbert (gilbertd  (at) indiana.edu) of the Department of Biology of the University of Indiana, has written SeqPup versions 0.9 and 0.6, a biological sequence editor and analysis program usable on MacOS, Windows and Unix systems. It allows alignment of sequences by hand and submission of selected parts of selected sequences to phylogeny programs, as well as to network services such as BLAST. It is available in a more complete earlier version (0.7) written in C++, or a later (0.9) version written in Java. The latter will work on all systems that have the Java 1.1 runtime environment. It will not work on the Java 1.2 runtime environment, so the earlier Java 1.1 environment needs to be installed as well if the later version of Java is present. (I do not know whether SeqPup works with later versions of Java). The two versions of SeqPup are described at a web page at http://iubio.bio.indiana.edu/soft/molbio/seqpup/java/seqpup-doc.html. Versions 0.9 and 0.6 can be obtained at its download web page The earlier version (0.6) is available in C++, and executables of it are available for MacOS (the PowerMac and 68K platforms), Windows, and Unix X windows systems including Sun Solaris, SGI Irix, Dec (HP/Compaq) Unix, Linux. The C++ source code for the 0.6 version is available by anonymous ftp at: iubio.bio.indiana.edu in directory util/dclap/source/.


  Matthias Wolf, Joachim Friedrich, Thomas Dandekar, and Tobias Müller of the Department of Bioinformatics at the the Biocenter, University of Würzburg, Germany (Matthias.Wolf (at) biozentrum.uni-wuerzburg.de) have written CBCAnalyzer versions 1.0.3 and 1.1β, programs for inferring phylogenies based on compensatory base changes (CBCs). The CBCAnalyzer (CBC = compensatory base change) is a custom written software toolbox consisting of three parts, CTTransform, CBCDetect, and CBCTree. CTTransform reads several CT-file formats (ct, RNAviz ct or Mac ct), and generates a so called "bracket-dot-bracket" format that specifies which sites are paired in an RNA structure. This typically is used as input for other tools such as RNAforester, RNAmovie or MARNA. The latter one creates a multiple alignment based on primary sequences and secondary structures that now can be used as input for CBCDetect. The count (distance) matrix of compensating changes obtained by CBCDetect is used as input for CBCTree that reconstructs a phylogram by using the BIONJ algorithm. It is described in the paper: Wolf, M., J. Friedrich, T. Dandekar and T. Müller. 2005. CBCAnalyzer: inferring phylogenies based on compensatory base changes in RNA secondary structures. In Silico Biology 5: 0027. It is available as C++ source code which can be compiled on Linux, and as Windows executables. It can be downloaded from its web site at http://cbcanalyzer.bioapps.biozentrum.uni-wuerzburg.de/


Wolfgang Ludwig and Oliver Strunk of the Lehrstuhl für Mikrobiologie of the Technische Universität München (wolfgang.ludwig  (at) arb-home.de) distribute ARB, an environment for 16s/18s/23s ribosomal RNA sequence data. It provides a windowing environment for building up databases of RNA sequences, aligning them, and searching, editing, modifying, aligning, profiling, and constructing trees. ARB uses its own RNA sequence databases which are made available to ARB over the Web. For phylogenies it uses programs from PHYLIP, AXML, PHYML and fastDNAml, as well as its own ARB Neighbor-Joining program. ARB is also incorporates a variety of other sequence analysis software. It can handle large numbers of sequences and has sophisticated tree drawing and manipulation. ARB is distributed as source code and executables for Linux systems, specifically Ubuntu and SuSE Linux. It can also be compiled successfully on SunOS and Mac OS X. ARB is available from its web site at http://www.arb-home.de/. A Mac OS X port of ARB is available by Mike Dyall-Smith of the Max Planck Institute of Biochemistry at http://www.haloarchaea.com/resources/arb/index.html.


Ian Holmes of the Department of Bioengineering of the University of California at Berkeley, Berkeley, Calfornia (ihh (at) berkeley.edu) has released DART (Dna, Amino and Rna Tests), a package of programs to do a variety of genomic, alignment, and RNA structure inferences. (DART is not to be confused with NCBI's Domain Architecture Retrieval Tool, also called DART). Various programs in the DART package do phylogenetic alignment; RNA structure prediction and multiple RNA alignment, stochastic context-free grammars, inference of evolutionary models, phylo-HMMs and phylo grammars, reconstruction of ancestral sequences; and phylogenomics. It contains a number of programs including

The programs use statistical algorithms (MCMC, EM) to impute multiple alignments, annotations and other unseen evolutionary parameters from sequence data. All are based on stochastic grammars or state-machine models of sequence mutation and natural selection. Many other people (listed at the DART web site) have contributed to the package. (The package includes “EvolDeeds”, which I have awarded the John Buettner-Janusch Award for most sinister name of a program). A large number of papers on the individual methods in the package are listed at the DART web site. It is available as C source code. It can be downloaded from its web site at http://biowiki.org/DART


Andrew Smith, Thomas W. H. Lui and Elisabeth Tillier of the Ontario Cancer Institute and the Department of Medical Biophysics at the University of Toronto, Canada (e.tillier (at) utoronto.ca) have released rRNA phylogeny, a program package to infer phylogenies from ribosomal RNA using a model of substitution that allows for compensating substiutions at paired sites. The program makes use of a model (the OTRNA model) of ribosomal RNA substitution that has different rates for paired and unpaired sites, that reflect the lower probability of a compensated substitution that maintains the pairing. The model is empirically tabulated from rRNA sequences and used in a modified version of programs from PHYLIP to infer phylogenies. The package also allows distances to be computed from the OTRNA model for use in distance matrix programs. The methods are described in the paper: Smith, A., T. W. H. Lui and E. R. M. Tillier, 2004. Empirical substitution models for Ribosomal RNA Molecular Biology and Evolution 21: 419-427. It is available as C source code, Windows executables and Linux executables. It can be downloaded from its web site at http://www.uhnresearch.ca/labs/tillier/software.htm


Dick Hwang and Phil Green of the Department of Genome Sciences of the University of Washington, Seattle, Washington (dhwang (at) u.washington.edu) has released AMBIORE (Applications for Mcmc Bayesian Inference Of Rates in Evolution), version 1.00, which estimates rates of change in nucleotide sequences depending on the two neighboring bases. It implements a flexible and computationally efficient Bayesian Markov chain Monte Carlo approach to estimating rates in evolution given a sequence alignment and tree topology relating the species. The evolutionary model allows substitution rates at a site to depend on the two flanking nucleotides, the branch of the phylogenetic tree, and position within a sequence. The methods are described in the paper: Hwang D. G. and P. Green. 2004. Bayesian Markov chain Monte Carlo sequence analysis reveals varying neutral substitution patterns in mammalian evolution. Proceedings of the National Academy of Sciences U.S.A. 101: 13994-14001. It is available as C source code. It can be downloaded from its web site at http://www.phrap.org/othersoftware.html


Chris Saunders, then of the Department of Genome Sciences of the University of Washington, Seattle, Washington (and currently at Illumina, Inc.) has written CodeAxe version 1.0, a tool for the phylogenetic analysis of mutation and selection in coding sequences. It provides, for a phylogeny provided by the user, analysis of approximate maximum likelihood nucleotide, codon and "extended codon" models that allow for neighbor-dependent nucleotide mutation, asymmetric exchange-dependent amino acid selection, nonreversible evolution and mixture models of mutation and selection classes. It is described in the paper: Saunders, C. T. and P. Green. 2007. Insights from modeling protein evolution with context-dependent mutation and asymmetric amino acid selection. Molecular Biology and Evolution 24(12): 2632-2647. It is available as C++ source code. It can be downloaded from its web site at http://www.phrap.org/othersoftware.html


[BioEdit icon] Tom Hall, then of the Department of Microbiology at North Carolina State University, and now of Ibis Therapeutics, Carlsbad, Calfornia (thall  (at) isisph.com) has produced BioEdit, version 7.0.5. This is a sequence editor with many kinds of general molecular biology functions available (alignment, BLAST searches, plasmid drawing, restriction mapping, sequence machine trace viewing, etc.). For our purposes the feature worth mentioning is that it comes with a number of existing phylogeny programs which can be automatically run from within BioEdit. These are: TreeView, fastDNAml, and six DNA and protein programs from PHYLIP. It has been described in the paper: Hall, T. A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Sympoisum Series 41: 95-98. BioEdit is available as Windows executables. Version 7.0.5 is available from its web site at http://www.mbio.ncsu.edu/BioEdit/bioedit.html.


  GeneStudio, Inc. of Suwanee, Georgia (info  (at) genestudio.com) has released GeneStudio Pro, version 2.1.2.4, a commercial package for sequence analysis and sequence format conversion. (Incidentally, although the Suwanee River is partly located in Georgia, the town of Suwanee is not located "way down upon the Suwanee River" but is quite far from it). Included is an Alignment Editor that can invoke a variety of phylogeny programs, including fastDNAml, some programs from PHYLIP, Tree-puzzle and TreeView. It can also do many other functions such as sequence format conversion, BLAST searches, and contig editing. GeneStudio Pro is for the Windows platform. A free trial version is available. Prices are not given on their web site but are available on request from a page on their web site, accessed through the trial version. For further information see the GeneStudio Pro web site at http://www.genestudio.com/genestudio.htm.


[Simplot
icon  Stuart Ray, of the Division of Infectious Diseases of the Department of Medicine at the Johns Hopkins University School of Medicine, Baltimore, Maryland (sray (at) jhmi.edu) has produced Simplot, version 3.5.1. It is a Windows program that serves as a front end to either code from PHYLIP or to PAUP* and enables you to easily submit jobs to them. SIMPLOT enables you to select regions and do other forms of data selection. It can also carry out the "bootscanning" method of detecting inconsistencies in trees in different regions of a sequence, which can be a signal for recombination. It was first described in the paper: Lole, K. S., R. C. Bollinger, R. S. Paranjape, D. Gadkari, S. S. Kulkarni, M. G. Novak, R. Ingersoll, H. W. Sheppard, and S. C. Ray. 1999. Full-length human immunodeficiency virus type 1 genomes from subtype C-infected seroconverters in India, with evidence of intersubtype recombination. Journal of Virology 73(1): 152-160. It is distributed as a Windows executable through its web site at http://sray.med.som.jhmi.edu/SCRoftware/simplot/


James J. Cai of the Department of the Department of Veterinary Integrative Biosciences at the Texas A&M University, College Station, Texas (jcai (at) tamu.edu) has written MBEToolbox (Molecular Biology and Evolution Toolbox), version 2.21, A MATLAB package to enable evolutionary biologists to analyze and view DNA and protein sequences. MBEToolbox includes sequence manipulation and statistics, evolutionary distance calculations, tree creation, a novel window analysis method and a graphical user interface. version 2.0 added new functions for phylogenetic analyses by the maximum likelihood method, analysis of site-specific evolutionary rates, and algorithms to detect recombination. It is described in the papers:

It is available as a MATLAB package, and has versions for download to Windows, Linux, and Sun Solaris systems. It requires MATLAB 6.5 or higher; the parsimony and likelihood analyses also require programs from PHYLIP. MBEToolbox can be downloaded from its web site at http://bioinformatics.org/mbetoolbox/


[Bosque icon]  Salvador Ramirez of the Departamento de Oceanografia of the Universidad de Concepcion, Chile (sram (at) profc.udec.cl) has released Bosque Phylogenetic Analysis Software, version 1.7.42, an integrated set of phylogenetic analysis software into a complete graphical user interface. Bosque is a graphical program that integrates console programs from the PHYLIP package, from TREE-PUZZLE, and from MUSCLE. As a graphical program it also includes sequence, alignment and tree editors to ease the manipulation of the data, the execution of the programs and the administration of the output result from the phylogenetic programs. It also has the option to function as a client-server program where a Bosque server is installed, thus allowing the remote execution of the mentioned phylogenetic programs. It is available as Windows executables, Mac OS X executables, and Linux executables. It can be downloaded from its web site at http://bosque.udec.cl


John Archer and David Robertson of the Faculty of Life Sciences of the University of Manchester (john.archer.jpa (at) googlemail.com) and Andrew Rambaut of the Institute of Evolutionary Biology of the University of Edinburgh (a.rambaut (at) ed.ac.uk) have produced Segminator II, a tool for extracting data from reads from viral data generated using the 454 Life Science sequencing platform. Following the mapping and pairwise alignment of reads, data that is immediately accessible in tabulated format includes:

  1. Obtaining nucleotide and amino acid residue frequencies at sites across the template.
  2. Generating a consensus across data set using overlapping windows (while allowing for indels differing from those in relation to the template).
  3. Removal of reads based on hamming distance cutoff thresholds.
  4. Reads spanning a particular region of the template can be multiply aligned and used to infer phylogenetic trees. and
  5. Identification of variants within viral populations.
The user can make a quick phylogenetic scan of clusters within the data using the TreeDar feature, or a detailed phylogenetic analysis of reads spanning a selected region of the template with the ability to track individual codon states on the tree. Segminator uses MUSCLE and RAxML (which the user must install separately) to produce maximum likelihood trees for the phylogeny step. Segminator is described in the paper: Archer, J., A. Rambaut, B. E. Taillon, R. Harrigan, M. Lewis and D. Robertson. 2010. The evolutionary analysis of emerging low frequency HIV-1 CXCR4-using variants through time - an ultra-deep approach. PLoS Computational Biology 6 (12): e1001022. It is available as Java executables with associated Windows executables, Linux executables and Mac OS X universal executables. It can be downloaded from its web site at http://www.bioinf.manchester.ac.uk/segminator/


[Likewind icon]   John Archibald and Andrew Roger, of the Department of Biochemistry and Molecular Biology, Dalhousie University, Halifax, Nova Scotia, Canada (john.archibald (at) dal.ca and aroger (at) is.dal.ca) have released Likewind version 1.0.1. This is a set of three Perl scripts that allow the user to create the necessary PAUP* blocks to drive a likelihood analysis of differences in phylogenies betweend different parts of a molecule. The object is to detect recombination, hybridization, or other sources of conflicting phylogenetic signal. The utilities also harvest the output from PAUP* and present it to the user as a log-likelihood difference plot. This shows the log-likelihood difference in the window of adjacent sites between a predefined tree and the best estimate from the local information. The scripts need Seq-Gen and PAUP* present in a Linux or Unix environment. The method has been described in a paper: Archibald, J. M. and A. J. Roger. 2002. Gene conversion and the evolution of euryarchaeal chaperonins: a maximum likelihood-based method for detecting conflicting phylogenetic signals. Journal of Molecular Evolution 55: 232-245. Likewind is available from its web site at http://rogerlab.biochemistryandmolecularbiology.dal.ca/Software/Software.htm#likewind


Christian Zmasek, currently of the Genomics Institute of the Novartis Research Foundation in San Diego, California (czmasek  (at) gnf.org) and Sean Eddy (eddy  (at)  genetics.wustl.edu) of the Department of Genetics, Washington University, St. Louis, Missouri have released FORESTER, version 1.92, a package of Java routines for inferring gene function using the output of high-throughput genome sequencing. It includes the RIO Resampled Inference of Orthologs method which searches for orthologs in the PFAM database, the SDI Speciation-Duplication Inference method, and ATV, a tree viewer. This can take a tree in the standard Newick format (and also in an extension that has additional information), and display it in various forms. These are described in papers:

The programs (in Java) are available from the Forester web site at http://www.phylosoft.org/forester/. A RIO web server is available from this group: see the listing in the list of servers in these pages.


Louxin Zhang (matzix  (at) nus.edu.sg) of the Department of Mathematics of the Faculty of Science, of the National University of Singapore has produced WebPHYLIP, version 2.0, a web interface for the PHYLIP package, which can submit jobs to it. It has been described in an article: Lim, A. and L. Zhang. 1999. WebPHYLIP: a web interface to PHYLIP. Bioinformatics 15(12): 1068-1069. There are versions as Windows executables and for Linux/Unix as C source code which can be compiled. It is available at this download page at http://www.math.nus.edu.sg/~matzlx/webphylip_download/ above address.


  Biomatters, Ltd. in Auckald, New Zealand (sales (at) geneious.com) has released Geneious version 5.3, a Java package for sequence management, searching, editing and phylogenies. Geneious is an integrated bioinformatics tool suite for manipulating, finding, sharing, and exploring biological data such as DNA sequences or proteins, phylogenies, 3D structure information, publications, etc. It features tree-based progressive sequence alignment and phylogenetic analysis by Neighbor-Joining and UPGMA, bootstapping, drawing the phylogeny, access to biological databases, BLAST, protein structure viewing, NCBI, EMBL, PubMed auto-find, and more. It includes an API for creating your own plugins. It is available as Java executables. Geneious is available for free; that version cannot edit sequences or annotations, cannot run the multiple alignment program ClustalW from within Geneious, and cannot do some other tasks. A full version, Geneious Pro, is available commercially. Free downloads of Geneious or purchases of Geneious Pro can be done from the Biomatters web site at http://www.biomatters.com. Geneious Pro is available from Biomatters, Ltd. for a 12-month license for academic or government institutions for $395, for a 12-month student license for $175, or for a 12-month commercial license for $995. The prices for perpetual licenses for these classes of users are $795, $395, and $1,995. Discount prices for multiuser licenses are also available. Additional plugins enabling it to run MrBayes, PAUP* (Geneious Pro only), PHYML, DualBrothers, and MAFFT are available free at the website.


Catherine Letondal of the Institute Pasteur, Paris, France (letondal (at) pasteur.fr) distributes PISE (Pasteur Institute Software Environment), a tool to generate Web interfaces for Molecular Biology programs. PISE allows the user to generate web page interfaces to many programs, including PHYLIP, EMBOSS, MOLPHY, and fastDNAML, Web interfaces for these are available in PISE. The web interfaces allow you to rapidly create a web server for running these programs. PISE is used to create the Pasteur Institute web server. PISE web servers allow the results of one program to the "piped" to another program. The web server generated is designed to be run on a Unix machine. It is described in the paper: Letondal C. 2001. A Web interface generator for molecular biology programs in Unix. Bioinformatics, 17(1): 73-82. It is available as Perl scripts. It can be downloaded from its web site at http://www.pasteur.fr/recherche/unites/sis/Pise/


MathWorks of Natick, Massachusetts have produced Bioinformatics Toolbox version 3.6, a MATLAB toolbox for bioinformatics. It has many functions for sequence analysis and microarray data, including multiple sequence alignment and consensus sequences. For this listing, the relevant ones are that it enables you to create and edit phylogenetic trees. You can calculate pairwise distances between aligned or unaligned nucleotide or amino acid sequences using a broad range of similarity metrics, such as Jukes-Cantor, p-distance, alignment-score, or a user-defined distance method. Phylogenetic trees are constructed using hierarchical linkage with a variety of techniques, including neighbor joining, single and complete linkage, and UPGMA. Bioinformatics Toolbox includes tools for weighting and rerooting trees, calculating subtrees, and calculating canonical forms of trees. Through the graphical user interface, you can prune, reorder, and rename branches; explore distances; and read or write Newick-formatted files. You can also use the annotation tools in MATLAB to create presentation-quality trees. It is available as a MATLAB package. It is available from MathWorks, Inc. The price is not available on the website without a login, but there are Commercial, Academic, and Student prices. It us believed that commercial licenses are about $1,000, academic licenses about $90. More information and sales contact can be obtained from the product web site at http://www.mathworks.com/products/bioinfo/


 The UGENE team of the Unipro company, Novosibirsk, Russia (mfursov (at) unipro.ru) has released UGENE version 1.8.0, a free cross-platform genome analysis suite. It is a sequence manipulation environment which can call other programs as plugins, including PHYLIP programs to compute distances and create trees from distance matrix methods, and MUSCLE to align sequences. It can also plot trees. It is available as C++ source code, Windows executables, Linux executables, Powermac Mac OS X executables and Intel Mac OS X executables. It can be downloaded from its web site at http://ugene.unipro.ru


   Andrew Rambaut of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) has written Phylogen version 1.1, a tree simulation program. It simulates phylogenies produced by a birth-death process. It also has a mode that allows for multiple epdisodes of evolution and multiple mass extinctions. It is available as generic source code for Unix and as Mac OS, Mac OS X, and Windows executables. It is distributed from its web page at http://tree.bio.ed.ac.uk/software/phylogen/. Although Rambaut considers it to have superseded his earlier program Bi-De. Phylogen is itself now unsupported software that will be distributed but not developed further.


[Genie icon here] Oliver Pybus, of the Department of Zoology, University of Oxford, (oliver.pybus  (at) zoo.ox.ac.uk and Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) have written Genie (GENealogical Interval Explorer) version 3.0, a program for the inference of demographic history from reconstructed phylogenies. The methods it implements are described in the papers:

It is considered by its authors to supersede the methods used in Rambaut's program End-Epi, which has been withdrawn from distribution. Genie is available from its web site at http://evolve.zoo.ox.ac.uk/evolve/Genie.html as Mac OS, Linux or Windows executables, and as C source code for Unix.


[MacroCAIC icon] Paul-Michael Agapow (p.agapow  (at) ic.ac.uk) and Nick Isaac, then of the Department of Biology, Imperial College, Silwood Park, U.K. (and most recently of the Health Protection Agency, U.K.) has released MacroCAIC version 1.0.1, which was developed from CAIC, by Andy Purvis and Andrew Rambaut. MacroCAIC uses phylogenies and data sets of character values to examine correlates of species richness, i.e. which traits are associated with abnormal speciation. The phylogeny is used to make these correlations independent. Traits may be continuous or discrete, and not every trait value needs to be known for every clade in the phylogeny. MacroCAIC has been described in a paper: Agapow, P.-M. and N. J. B. Isaac. 2002. MacroCAIC: correlates of species richness. Diversity & Distributions 8: 41-43. MacroCAIC is a PowerMac and 68k Mac binary executable for Mac OS. It is available from its web site at http://www.bio.ic.ac.uk/evolve/software/macrocaic/index.html.


[Seq-Gen icon here]   Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) University of Oxford, (andrew.rambaut  (at) zoo.ox.ac.uk) and Nick Grassly of the Department of Infectious Disease Epidemiology of Imperial College School of Medicine, St. Mary's Campus, London (n.grassly  (at) ic.ac.uk) have written Seq-Gen (Sequence Generator), version 1.3.2, a program that will simulate the evolution of nucleotide sequences and protein sequences along a phylogeny or multiple phylogenies, using common models of the nucleotide or protein substitution process. A range of models of molecular evolution are implemented Nucleotide frequencies and other parameters of the model may be given and site-specific rate heterogeneity may also be incorporated in a number of ways. The models available are the Hasegawa, Kishino and Yano (HKY) model, the Felsenstein F84 model, the general reversible model, the Kimura 2-parameter model, the Jukes-Cantor model and the Dayhoff PAM, JTT, Blosum62, mtREV, and WAG amino acid substitution models. Rate heterogeneity among sites or among the different positions within a codon can be specified. Seq-Gen is described in a paper: Rambaut, A. and N. C. Grassly. 1997. Seq-Gen: an application for the Monte Carlo simulation of DNA sequence evolution along phylogenetic trees. Computer Applications in the Biosciences ICABIOS) 13: 235-238. A Mac OS executable is available, as well as source code files for Unix systems. A Windows executable is available for an earlier version, 1.3.1. These are available from its Web page at http://tree.bio.ed.ac.uk/software/seqgen/.


  Tom Wilcox has written SG Runner version 2.0.1, a graphical user interface for running Seq-Gen. It is available as Powermac Mac OS X executables. It can be downloaded from his software web site at http://homepage.mac.com/tpwilcox/FileSharing15.html


[PSeq-Gen icon here] Nick Grassly of the Department of Infectious Disease Epidemiology of Imperial College School of Medicine, St. Mary's Campus, London (n.grassly  (at) ic.ac.uk) and Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) have written PSeq-Gen (Protein-Sequence Generator), version 1.1, which will simulate the evolution of protein sequences along evolutionary trees. Three common models of amino acid substitution are implemented (PAM, JTT, and mtREV), allow for user-defined amino acid frequencies. Site-specific rate heterogeneity following a gamma distribution is allowed. The program can handle multiple trees and produce multiple data sets. PSeq-Gen is available from a web site at the IUBIO server at http://iubio.bio.indiana.edu/soft/iubionew/molbio/evolution/phylo/PSeq-Gen/main.html as Unix source code and also as Mac OS executables. An online manual can also be viewed at that site. PSeq-Gen is now largely superseded by Seq-Gen, and although it is still available at the IUBIO server it is not being actively supported by its authors.


[Treevolve icon here] Nick Grassly, of the School of Public Health of Imperial College, St. Mary's Campus, London (n.grassly (at) imperial.ac.uk) and Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) wrote Treevolve, version 1.32 and also Ptreevolve, programs that simulate the evolution of DNA and protein sequences respectively. The molecular sequences are simulated under coalescent models with constant population size, or with exponential population size growth. In addition different levels of recombination can be specified. In Treevolve, it is also possible to have an island model of population subdivision. Treevolve and Ptreevolve are written in ANSI C and should compile on most Unix systems. They are also available as Mac OS executables, and a project file for the MetroWerks Codewarrior compiler is included in the Macintosh archive. They can be obtained from programs viewed, from their archives at the IUBIO web site at http://microbe.bio.indiana.edu:7131/soft/iubionew/molbio/genetic/pop/TreeVolve/


Jens Stoye, Dirk Evers and Folker Meyer of the Research Center for Interdisciplinary Studies on Structure Formation (FSPM) and the Technische Fakultät of the University of Bielefeld, Germany (j.stoye  (at) dkfz-heidelberg.de, dirk  (at)  TechFak.Uni-Bielefeld.de, and folker  (at)  TechFak.Uni-Bielefeld.de) have released ROSE, the Random model Of Sequence Evolution, version 1.3. It simulates the evolution of DNA, RNA, or protein sequences on a randomly generated tree, allowing insertions and deletions and substitution at different rates at different sites as well. It can also use a predefined tree that is input in standard format. It can report ancestral sequences or sequences at the tips of the tree, and it also keeps a record of the true multiple sequence alignment for comparison with the results of multiple sequence alignment programs. ROSE is described in the paper: Stoye, J. D. Evers and F. Meyer. 1998. Rose: generating sequence families. Bioinformatics 14: 157-163. ROSE is available in source code at its web site at http://bibiserv.techfak.uni-bielefeld.de/rose/.


Andy Pang, Andrew Smith, Paulo Nuin and Elisabeth Tillier of the Cancer Genomics and Proteomics Division of the Ontario Cancer Institute and the Department of Medical Biophysics of the University of Toronto, Canada (e.tillier (at) utoronto.ca) have released Simprot (Simulation of Proteins), version 1.03, a program to simulate protein evolution by substitution, insertion and deletion. It allows for several models of amino acid substitution (PAM, JTT and PMB), it allows for gamma distributed sites rates according to Yang's model, and it implements a parameterised Qian and Goldstein distribution model for insertion and deletion. It is described in the paper: Pang, A., A. Smith., P. Nuin, and E. R. M. Tillier. 2005. SIMPROT: Using an empirically determined indel distribution in simulations of protein evolution BMC Bioinformatics 6: 236. It is available as C source code and Windows executables. It can be downloaded from its web site at http://www.uhnresearch.ca/labs/tillier/software.htm


Cory Strope, S. D. Scott and Etsuko Moriyama then of the Department of Computer Science and School of Biological Sciences of the University of Nebraska (corystrope (at) gmail.com) have released indel-Seq-Gen version 2.1.0, a sequence family simulator incorporating domains, motifs, and indels. it simulates a more realistic evolutionary process of protein sequences including insertions and deletions (indels). iSG allows the user to simulate multiple subsequences according to different evolutionary parameters, which is necessary for generating realistic protein families with multiple domains. It tracks all evolutionary events including indels and outputs the "true" multiple alignment of the simulated sequences. iSG can also generate a larger sequence space by allowing the use of multiple related root sequences. It is intended to be used to test the accuracy of multiple alignment methods, phylogenetic methods, evolutionary hypotheses, ancestral protein reconstruction methods, and protein family classification methods. It is described in the paper: Strope, C. L., S. D. Scott, and E. N. Moriyama. 2007. indel-Seq-Gen: A new protein family simulator incorporating domains, motifs, and indels. Molecular Biology and Evolution 24: 640-649. It is available as C source code, Perl script, Linux executables and Intel Mac OS X executables. It can be downloaded from its web site at http://bioinfolab.unl.edu/~cstrope/iSG


[ProSeq icon]  Dmitry Filatov, of the Department of Plant Sciences of the University of Oxford, U.K. (dmitry.filatov  (at) plants.ox.ac.uk) has released ProSeq (Processor of Sequences) version 3. ProSeq is a sequence-editing environment that can do sequence alignment editing, translation, detection of polymorphic sites, and a variety of tests, many of a population-genetic nature, for neutrality and recombination. The part of its capabilities that are relevant to this listing is that it can simulate the evolution of a set of DNA sequences along a coalescent tree, with or without recombination. Version 3 allows simultaneous alignment of multiple loci. An earlier version of ProSeq is described in a paper: Filatov, D.A. 2002. ProSeq: A software for preparation and evolutionary analysis of DNA sequence data sets. Molecular Ecology Notes 2: 621-624. ProSeq is available as Windows or Linux executables from its web site at http://dps.plants.ox.ac.uk/sequencing/proseq.htm.


  Paul Michael Agapow (agapow (at) agapow.net), then of the Department of Biology, Imperial College, Silwood Park, U.K. (most recently of the Health Protection Agency, U.K.) has released MESA (MacroEvolutionary analysis and SimulAtion), version 1.9.23, a program to simulate evolution of a group, allowing for a variety of kinds of extinction mechanisms. It can simulate evolution and describe the diversity of the resulting groups. Its actions can be recorded and repeated on multiple trees. It also has features for editing and manipulating trees. MESA is available as an executable for Windows and as an executable for Mac OS X from its web site at http://www.agapow.net/software/mesa


[DAWG icon]  Reed Cartwright of the Department of Biology and Biochemistry at the University of Houston, Houston, Texas (reed (at) scit.us) has written DAWG (DNA Assembly With Gaps), version 1.2, a program to simulate evolution of DNA sequences with recombination and gaps. It is designed to simulate the evolution of recombinant DNA sequences in continuous time based on the general time reversible model with gamma and invariant rate heterogeneity and a novel length-dependent model of gap formation. It accepts phylogenies in Newick format and can return the sequence of any node, allowing for the exact evolutionary history to be recorded at the discretion of users. Dawg records the gap history of every lineage to produce the true alignment in the output. Many options are available to allow users to customize their simulations and results. It is described in the paper: Cartwright, R. 2005. DNA Assembly with gaps (DAWG): simulating sequence evolution. Bioinformatics 21 (Supplement 3): iii31-iii38. It is available as C source code. It can be downloaded from its web site at http://scit.us/projects/dawg/wiki


Barry Hall of the Bellingham Research Institute in Bellingham, Washington (barryhall (at) zeninternet.com) has written EvolveAGene3, a program that simulates evolution of a protein sequence along a tree. It generates a bifurcating tree, and assigns branch lengths from a distribution whose mean is specified by the user. A protein sequence is evolved along this tree, with deletions and insertions of codons and with base subtitutions. Substitutions that change the amino acid are accepted with a specified probability. This includes having variable regions of selection intensity, positive or purifying, within the sequence and variation in intensity of selection over branches. Output includes the true tree and unaligned coding sequences and protein sequences as well as the true DNA and the true protein alignments. It is available as Perl script, Windows executables and Mac OS X universal executables. See the web site http://web.me.com/barryghall/Software/Software.html


[EvolSimulator icon]   Robert Beiko and Robert Charlebois of the Faculty of Computer Science at Dalhousie University (beiko (at) cs.dal.ca) have released EvolSimulator version 2.1.0, a program to simulate the evolution of genes and genomes. EvolSimulator was created to allow the simulation of complex evolutionary regimes, potentially comprising nonstationary and nonuniform processes at the sequence level, alongside genome-level processes such as gene duplication, gene loss, and lateral gene transfer. Several models of LGT are implemented, including random transfers, transfers that favour close relatives in the organismal tree, and preferential transfer among members of the same habitat. It is described in the paper: Beiko, R. G. and R. L. Charlebois. 2007. A simulation test bed for hypotheses of genome evolution. Bioinformatics 23 (7): 825-31. It is available as C++ source code. It can be downloaded from its web site at http://bioinformatics.org.au/evolsim


[Recodon icon]   Miguel Arenas and David Posada of the Department of Biochemistry, Genetics and Immunology of the University of Vigo, Spain (miguelmmmab (at) gmail.com) have released Recodon (coalescent simulation of Coding sequences with Recombination), version 1.6.0, A simulator of coding sequences based on the coalescent with recombination, migration and demographics. The program simulates coding and non-coding sequences using the coalescent and with recombination, migration and demographics. It implements all major DNA substitution models of evolution and the GY94 codon model including rate heterogenity among codons. It is described in the paper: Arenas, M. and D. Posada, 2007. Recodon: Coalescent simulation of coding DNA sequences with recombination, migration and demography. BMC Bioinformatics 8: 458. It is available as C source code, Windows executables, Linux executables and Powermac Mac OS X executables. It can be downloaded from its web site at http://darwin.uvigo.es/software/recodon.html


[NetRecodon icon]  Miguel Arenas and David Posada of the Department of Biochemistry, Genetics and Immunology of the University of Vigo, Spain (miguelmmmab (at) gmail.com) have released NetRecodon (coalescent simulation with intracodon Recombination), version 6.0.0, A simulator of coding sequences based on the coalescent with (intra and inter codon) recombination, migration and demographics. The program simulates coding and non-coding sequences using the coalescent with recombination (where breakpoints can occur at both intra and inter codon positions), migration and demographics. Indeed, the program implements many DNA substitution models of evolution and also GY94 codon models including heterogenity of rate of evolution among codons. This program is more advanced than the previous program Recodon. One improvement is that it allows recombination within as well as between codons. It is described in the paper: Arenas, M. and D. Posada. 2010. Coalescent simulation of intracodon recombination. Genetics 184(2): 429-437. It is available as C source code, Windows executables, Linux executables and Powermac Mac OS X executables. It can be downloaded from its web site at http://darwin.uvigo.es/software/netrecodon.html


Martin Senger (senger  (at) ebi.ac.uk) of the European Bioinformatics Institute, Hinxton, U.K. and Peter Ernst (P.Ernst  (at) dkfz-heidelberg.de) of the Deutsches Krebsforschungszentrum, Heidelberg, Germany distribute W2H, (Web 2 Husar) version 4.1.2, a web interface for running molecular sequence analysis programs. It can invoke a variety of sequence analysis programs, including the EMBOSS Embassy versions of a number of PHYLIP 3.5 programs. W2H is a set of web pages that has been developed primarily with Unix platforms in mind. They can be downloaded and are available from the W2H web page at http://www.w2h.dkfz-heidelberg.de/


  Juan José de Haro (jjdeharo (at) gmail.com) of "una pequeña ciudad" of the province of Barcelona, Spain, has released Phyledit version 2.0. This is an interactive data editing and analysis program that uses PHYLIP and Treeview for analysis and display of trees. It uses eight programs from PHYLIP, all concerned with discrete 0/1 characters. Phyledit's menus, responses and documentation are all in Spanish. It is a Windows executable downloadable available from its web site at http://perso.wanadoo.es/jjdeharo/phyledit/index.htm. It is available there either by itself or with the PHYLIP and TREEVIEW executables as well.


  Øyvind Hammer, of the Paleontological Museum of the University of Oslo, Norway (ohammer  (at) nhm.uio.no), together with David A.T. Harper of the Geological Museum, Copenhagen, Denmark, and P.D. Ryan, has written PAST (PAleontological STatistics), version 2.04, a package which carries out many kinds of paleontological data analyses, including stratigraphic and morphometric statistics. It also does parsimony analysis, including exhaustive, branch-and-bound and heuristic algorithms for Wagner, Fitch and Dollo parsimony. It does bootstrap methods, strict and majority rule consensus trees, and consistency and retention indices. It calculates three stratigraphic congruency indices with permutation tests. It also does many other statistics and curve fitting. PAST is described in a paper: Hammer, Ø., D.A.T. Harper, and P. D. Ryan. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4:, issue 1: http://palaeo-electronica.org/2001_1/past/issue1_01.htm. PAST is available from its web site at http://folk.uio.no/ohammer/past/index.html as a Windows executable. Manuals can be read online or downloaded from the web site.


[GHOSTS icon]    Matthew Wills of the Department of Biology at the the University of Bath,. U.K. (bssmaw (at) bath.ac.uk) has produced GHOSTS version 2.4, a program for significance tests for RCI, SCI and GER values by randomization. It calculates the Relative Completeness Index (RCI), the Stratigraphic Consistency Index (SCI), and the Gap Excess Ratio (GER) measures of consistency of phylogenies with the stratigraphic record if given one or more tree topologies and stratigraphic range data for up to 74 terminal taxa. It can be used with NEXUS tree files and can be used interactively with MacClade. It can also randomly permute the assignment of stratigraphic ranges among taxa, while holding tree topologies constant, to yield a distribution of values. It tests whether the RCI and SCI values for the original data differ significantly from the random distributions. Its methods are described in the paper: Wills, M. A. 1999. Congruence between phylogeny and stratigraphy: randomization tests. Systematic Biology 48 559-580. It is available as source code for use with Chipmunk Basic. It can be downloaded from the Biodiversity Lab resources web page at http://www.bath.ac.uk/bio-sci/biodiversity-lab/?page_id=281  The program requires a password that can then be obtained from him.


[QDate icon here] Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) has written QDate version 1.1.1. QDate estimates the date of divergence between two pairs of sequences given that the date of divergence of the members of each pairs is known. It analyzes the data under three models: (1) a perfectly clocklike model, (2) a model in which one pair has a different rate of divergence than the other, and (3) a model in which all branches have different rates. The method is described in the paper: Rambaut, A., and L. Bromham. 1998. Estimating divergence dates from molecular sequences. Molecular Biology and Evolution 15: 442-448. QDate is available from its web page at the IUBIO server at ftp://net.bio.net/iubionew/molbio/evolution/phylo/QDate/main.html. It is available as C source code for Unix or as a Macintosh or Windows executable.


Marc Robinson-Rechavi, of the Department of Ecology and Evolution of the Université de Lausanne, Lausanne, Switzerland (Marc.Robinson-Rechavi  (at) unil.ch) has written RRTree, (Relative Rate tests within a Tree), version 1.1. It carries out relative rate tests for equality of evolutionary rates in DNA or protein sequences between lineages, taking into account the structure of the tree, which can be input in a number of common formats. In addition sequences are read in. The methods are described in a papers:

Marc Robinson-Rechavi suggests that other programs might do better and maintains the source code and documentation in distribution in case they might be of interest, but the program is not being further developed. RRtree is available as C source code and documentation from its web page at http://bioinfo.unil.ch/rrtree/


Bette Korber of the Theoretical Division, Los Alamos National Laboratory , Los Alamos, New Mexico (btk  (at) lanl.gov), Mark Muldoon and and their colleagues have released programs for rate evolution which implement the method described by Jeff Thorne, Hirohisa Kishino, and Ian Painter of North Carolina State University for making a maximum likelihood estimate of the phylogeny while allowing rates of evolution to change along the tree in a "relaxed molecular clock". Their method was described in a paper: Thorne, J. L., H. Kishino, and I. S. Painter. 1998. Estimating the rate of evolution of the rate of molecular evolution. Molecular Biology and Evolution 15: 1647-1657. Korber and her colleagues used it for their paper: Korber, B., M. Muldoon, J. Theiler, F. Gao, R. Gupta, A. Lapedes, B. H. Hahn, S. Wolinksy and T. Bhattacharya. 2000. Timing the ancestor of the HIV-1 pandemic strains. Science 288: 1789-1796. Three programs are available (all as C code for Unix):

The programs are available from the web site for the programs from that paper at http://www.santafe.edu/~btk/science-paper/bette.html. Two other programs used for the analyses in the paper are available there too.


Jeff Thorne of the Department of Genetics at North Carolina State University, Raleigh, North Carolina (thorne (at) statgen.ncsu.edu) has written Multidivtime, version 09.25.03, a program that estimates divergence times from multiple-locus data. The method starts with a given tree supplied by the user. The Bayesian dating method implemented in multidivtime uses a probabilistic model to describe the change in evolutionary rate over time and uses a Markov chain Monte Carlo (MCMC) procedure to derive the posterior distribution of rates and times. It allows simultaneous use of different substitution models for multiple data partitions as well as multiple calibration windows, and provides direct credibility intervals for estimated divergence times and substitution rates. The dating method is described in an earlier paper: Thorne, J. L., H. Kishino, and I. S. Painter. 1998. Estimating the rate of evolution of the rate of molecular evolution. Molecular Biology and Evolution 15: 1647-1657. It is available as C source code which can be compiled on Mac OS X or other standard Unix systems, and as Windows executables. Multidivtime is to be used in combination with some of the programs from Ziheng Yang's PAML package. Multidivtime can be downloaded from its web site at http://statgen.ncsu.edu/thorne/multidivtime.html. An earlier version is available from Ziheng Yang's lab at their web site at http://abacus.gene.ucl.ac.uk/software.html where it is called T3, the Thornian Time Traveler. Yang suggests using the more recent version at Thorne's site.


Tae-Kun Seo of the Professional Programme for Agricultural Bioinformatics of the University of Tokyo, Japan (seo (at) iu.a.u-tokyo.ac.jp) has written CodonRates version 1.0, a program to estimate divergence times and absolute rates of synonymous and nonsynonymous substitution with protein-coding genes. Given sequences and a tree, CodonRates can separate the amount of evolution into time and the synonymous and nonsynonymous rates without using a molecular clock. Data other than sequences, such as fossil constraints and a prior distribution for the root time, are required. It is described in the paper: Seo T-K, H. Kishino, and J. L. Thorne. 2004. Estimating absolute rates of synonymous and nonsynonymous nucleotide substitution in order to characterize natural selection and date species divergences. Molecular Biology and Evolution 21(7): 1201-1213. It is available as C++ source code and Windows executables. It can be downloaded from its web site at http://www.iu.a.u-tokyo.ac.jp/~seo/software.htm


Tom Britton, Cajsa Anderson, David Jacquet, Samuel Lundqvist, and Kåre Bremer of the Department of Mathematics and the Department of Systematic Botany of Stockholm University, Stockholm, Sweden (Anderson is at Uppsala Univerity) (tomb (at) math.su.se) have released PATHd8 (Path-date), a program to estimate divergence times from a nonclocklike tree and time constraints. PATHd8 reads in a tree which has branch lengths but with no requirement that these show a molecular clock. It also reads in constraints on the divergence times. These are then reconciled using either the PATHd8 method or the Mean Path Length method. It is described in the papers:

It is available as C source code, Windows executables, Linux executables, Powermac Mac OS X executables, and Sun SPARC executables. It can be downloaded from its web site at http://www.math.su.se/PATHd8/index.html


Bette Korber of the Theoretical Division, Los Alamos National Laboratory , Los Alamos, New Mexico (btk  (at) t10.lanl.gov), and her colleagues have released BRANCHLENGTH, a pair of programs which allow trees to have their branch lengths to any node plotted against time. They take output files from the PHYLIP program DNAML, or from Gary Olsen's program fastDNAmlM, or from one of the modified versions of fastDNAml as input. One program (BRANCHLENGTHxTIME) plots branch lengths to any specified node against time, makes best fit lines, and does Monte Carlo resampling statistics. The other program (BRANCHLENGTH) computes The sum of branch lengths to any specified node. Korber and her colleagues used it for their paper: Korber, B., M. Muldoon, J. Theiler, F. Gao, R. Gupta, A. Lapedes, B. H. Hahn, S. Wolinksy and T. Bhattacharya. 2000. Timing the ancestor of the HIV-1 pandemic strains. Science 288: 1789-1796. The programs are available from the web site for the programs from that paper at http://www.santafe.edu/~btk/science-paper/bette.html. Some further code (scripts and C code) to fully implement the methods used for branch length plotting in that paper is available from Korber on request to individuals who have the book Numerical Recipes and thus have a license to use code derived from it.


Kai Müller of the Institute for Evolution and Biodiversity of the Westfälische Wilhelms-Universität Münster, Münster, Germany (kaimueller (at) uni-munster.de) has written GRate (Group relative RATE tests), version 1.0, a program for carrying out relative rate tests of evolution in specified groups. GRate runs PAUP* and uses the sitewise log likelihoods and bootstrapping that it makes available, and analyzes these to carry out relative rate tests that compare the rate of evolution in two or more clades selected by the user. It is available as Windows executables. It can be downloaded from its web site at http://bioinfweb.info/Software/GRate


Nicolas Galtier and Manolo Gouy of the Institut des Sciences de l' Evolution of the Université Montpellier 2 - CNRS, Montpellier, France and the Laboratoire de Biométrie et Biologie Evolutive of the Universiteacute; Claude Bernard - Lyon, France (galtier (at) univ-montp2.fr) have produced NHML (NonHomogenous Maximum Likelihood), version 3, a maximum likelihood phylogeny program using a nonhomogenous model of nucleotide change. It uses a model developed by them which is not reversible and thus allows information about the location of the root of the tree. The program can evaluate a user-defined tree, or can start with a "star tree" and then resolve it by Subtree Pruning and Regrafting rearrangements to infer the phylogeny. It can use either a model of rates varying among sites or covarion-like model which allows rates at each site to independently vary along the branches of the tree. The methods are described in the papers:

NHML is available as C source code and Linux executables. It can be downloaded from its web site at http://162.38.181.1/nhml/nhml.html


Mike Maciukenas, then at the Department of Microbiology of the University of Illinois, wrote a wonderful X-windows based interactive tree-plotting program called TreeTool, version 2.02. It takes as input a PHYLIP tree file, with branch lengths if they are provided, displays the tree in either rooted or unrooted form on any X-windows screen, and allows the user to modify the form of the tree and the placement of nodes and labels. When the tree is in final form the user can have it written to a Postscript or PICT file and/or printed to a Postscript-compatible printer. TreeTool is free as a C program for X windows using the Xview library. Xview seems to be available mostly on Sun workstations (it can also be installed on Linux). TreeTool is available It will be found in the IUBIO software archive at http://iubio.bio.indiana.edu/soft/molbio/unix/treetool. It is also included in the GDE sequence analysis environment mentioned. A Debian Linux Treetool package for Intel-compatible processors is available at its Debian web page at http://packages.debian.org/lenny/treetool. It has links to the Debian package for Xview.


[TreeView icon here]  [TreeView X icon here]   Rod Page of the Division of Environmental and Evolutionary Biology of the University of Glasgow, Scotland (r.page  (at) bio.gla.ac.uk), has written TreeView version 1.6.6, a program for displaying and manipulating trees. It can draw rooted and unrooted trees, display bootstrap values, and edit trees by moving branches, collapsing them, and rerooting. The program reads NEXUS, PHYLIP, and Hennig86 style tree files (including files produced by fastDNAml and CLUSTALW), and can save trees in the same formats so that it can convert trees among these formats. The Mac and Windows versions have almost identical interfaces. They can support the standard fonts available on Macs and PCs, and they support the standard PICT and Windows Metafile formats for output, allowing tree pictures to be copied into other applications, as well as being saved in files. There is a print preview and drag-and-drop facilities. Currently TreeView can read trees with up to 1000 taxa. The program is free, and can be obtained from its web page from http://taxonomy.zoology.gla.ac.uk/rod/treeview.html. It comes in Mac OS and Windows executable versions. There is also a Unix/Linux version, TreeView X, which does not yet implement all features of the program. Source code is available at its page at Google Code at http://code.google.com/p/treeviewx/its page at Google Code. There is also online help including an online manual.


[NJplot icon]  Manolo Gouy of the Laboratoire de Biomètrie et Biologie Evolutive of the University of Lyon, France (mgouy  (at) biomserv.univ-lyon1.fr), has produced NJplot, version 2.3-2, which plots rooted phylogenies (input in the standard form) and saves the plots as Postscript (for Mac OS, PICT) files. It displays branch lengths and bootstrap information (if present) and allows the user to swap branches and change the position of the root. It is described in the paper Perrière, G. and M. Gouy 1996. WWW-Query: an on-line retrieval system for biological sequence banks. Biochimie 78: 364-369. It is available free as executables for Mac OS X, Mac OS, Windows, SunOS, Sun Solaris, Linux and DEC/Compaq/HP Alpha. It can be retrieved using its web page at http://pbil.univ-lyon1.fr/software/njplot.html. A Debian Linux package is available from its web page at http://packages.debian.org/unstable/science/njplot. It has links to a number of other Debian packages that are needed to run it, including Lesstif and the NCBI Vibrant toolkit. It is also available in a server at the Institut Pasteur in Paris.


[unrooted icon]  Manolo Gouy of the Laboratoire de Biomètrie et Biologie Evolutive of the University of Lyon, France (mgouy  (at) biomserv.univ-lyon1.fr), has written unrooted, which draws unrooted phylogenies and saves them in Postscript files (for Macintosh, PICT files). It is available free as executables for Mac OS X, Mac OS, Windows, SunOS, Sun Solaris, SGI, IBM Unix, Linux and DEC/Compaq/HP Alpha. It can be retrieved using its web page at http://pbil.univ-lyon1.fr/software/unrooted.html.


[DendroMaker icon]  Tadashi Imanishi (imanishi  (at) jbirc.aist.go.jp) of the Biological Information Research Center (BIRC) of the National Institute of Advanced Industrial Science and Technology (AIST), Tokyo, Japan has produced DendroMaker, version 4.1, Mac OS programs which can draw trees on the screen and print them in Postscript files or MacPaint files. They read the trees in from tree files produced by the Neighbor-Joining or UPGMA options of the Oden package, and can also read standard Newick-format tree files. They can them edit them and reroot them in a variety of ways. They can produce PICT files of the trees. DendroMaker is distributed as Mac OS PowerMac or 68k Mac executables (each available in both English and Japanese versions) from its web page at http://www.cib.nig.ac.jp/dda/timanish/dendromaker/home.html.


[Tree Draw Deck icon]  Don Gilbert (gilbertd  (at) bio.indiana.edu) of the Department of Biology of the University of Indiana, Bloomington, Indiana, has written Tree Draw Deck, a Hypercard deck that draws trees on the screen from standard Newick-format tree files. The deck, which runs on Macintoshes or PowerMacs that have Hypercard Player, is based on two programs, Drawtree and Drawgram, from the 3.3 version of PHYLIP. It allows mouse-driven interactive display of the trees with selection of options for display, and allows the resulting plot to to cut into the system clipboard and pasted into drawing programs such as MacDraw or Canvas. Its printing and file-saving options do not at present work, but the clipboard method works. It is available by anonymous ftp from ftp.bio.indiana.edu in directory molbio/mac. It is also available by ftp from ftp.ebi.ac.uk in directory pub/software/mac.


[Phylodendron icon]  Don Gilbert (gilbertd  (at) bio.indiana.edu) of the Department of Biology of the University of Indiana, Bloomington, Indiana, has written Phylodendron version 0.8d beta, a Java application for drawing phylogenetic trees. It will read tree data in standard Newick format, then display graphical views of the phylogenetic tree. Various options allow you to modify, adorn and edit the tree. Standard application functions to save, print, edit and manage preferences are included. This program will not estimate nor produce the tree data. Phylodendron is written as a Java application. It is also distributed with some extra parts for Mac OS. Phylodendron is an enhancement of the Mac Hypercard program Tree Draw Deck which was released by Gilbert in 1990, and uses tree drawing algorithms from PHYLIP. Phylodendron is available from its Web site http://iubio.bio.indiana.edu/soft/molbio/java/apps/trees/ or by anonymous ftp from iubio.bio.indiana.edu in directory molbio/java/apps/trees/. Source code is also available there. Phylodendron is also made available as a web server.


Julian Gough (gough  (at) cs.bris.ac.uk) of the Department of Computer Science at the University of Bristol, Bristol, U.K. has produced a Phylogenetic Tree Drawing Perl module that reads a tree and displays a rooted tree diagram. The user can specify width of the diagram and its line thickness, and can make each line consist of two shadings of different width, and add extra labels to tips. The routine is in Perl and is available at its web site at http://www.cs.bris.ac.uk/~gough/software/. Selecting the "Perl module" link there displays the Perl source on the browser, and you can then save it to disk. It is also available at his previous web site for it at http://supfam.mrc-lmb.cam.ac.uk/treedraw/tree.html


  Tamara Munzer (tmm  (at) cs.ubc.ca) and her colleagues at the Graphics, Visualization and Human-Compuyer-Interface group at the Department of Computer Science of the University of British Columbia, Vancouver, Canada have produced TreeJuxtaposer version 2.1. This displays a tree or shows two trees, with the locations of differences highlighted. Parts of the trees collapse and expand as one moves through the trees, so that very large trees can be examined. TreeJuxtaposer is a Java application which works with GL4Java, the Java implementation of the OpenGL graphics environment. TreeJuxtaposer is described in a paper for the SIGGRAPH 2003 conference: Munzner, T., F. Guimbretiere, S. Tasiran, L. Zhang, and Y. Zhou. 2003. TreeJuxtaposer: Scalable tree comparison using focus+context with guaranteed visibility. ACM Transactions on Graphics 22: 453-462. It was an overall winner in the IEEE Symposium on Information Visualization InfoVis 2003 Contest on "Visualization and Pair Wise Comparison of Trees". It can be downloaded from its web site at http://olduvai.sourceforge.net/tj/index.shtml


Christian Zmasek, currently of the Genomics Institute of the Novartis Research Foundation in San Diego, California (phylosoft  (at) gmail.com) has written Archaeopteryx, version 0.971β9m, a software tool for the visualization, analysis, and editing of potentially large and highly annotated phylogenetic trees. It is the successor to ATV, a tree viewer that is part of the FORESTER package. Archaeopteryx can read trees from a variety of tree file formats. It can do a number of different shapes of tree display, can have support values on the tree, allows different colors and display of various kinds of data such as photographs of the organisms. It can also display internal node data by mouse rollover as you move the mouse around the tree. Archaeopteryx is written in Java, and can be run on Sun Java version 5 and later. That version of Java is already installed in recent Windows and Mac OS X systems, and can also be installed by the user on Linux systems. Archaeopteryx is available at its web site at http://www.phylosoft.org/archaeopteryx/. There are also available there versions that can also be installed as an applet on Java-enabled browsers.


[Mavric]  Richard H. Ree of the Botany Department of the Field Museum of Natural History, Chicago, Illinois (rree  (at) fieldmuseum.org) has written Mavric (recursively, MAvric Visualizes RIck's Cladograms), version 0.8.3, a package in the Python language for the manipulation and visualization of phylogenetic data. It is intended to be flexible and easily customizable to suit the needs of phylogenetic biologists who use python. Mavric is named for its core application Mavric, which aims to provide a graphical interface to phylogenies in a manner similar to MacClade. (Rick notes that “the recursion that defines the name Mavric ultimately boils down to its central application: a tool to Manipulate And Visualize RIck's Cladograms -- if you followed all this you're pretty far ahead in understanding recursion”). Currently, its main usefulness is to view and manipulate phylogenetic trees. Branches can be moved around, pruned, rotated, have lengths changed. Trees can be grafted together, and viewed either rooted or unrooted. Postscript tree plots can be produced. Mavric is available as Python source code from its web site at http://www.bioinformatics.org/mavric/index.html.


[TreeEdit icon]  Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) and Mike Charleston (mcharles  (at) it.usyd.edu.au) of the Sydney University Biological Informatics and Technology Centre, Sydney, Australia have produced the TreeEdit Phylogenetic Tree Editor version 1.0a10. TreeEdit is an application for organizing, manipulating and viewing sets of trees. It is intended as a tool for preparing sets of trees for use in phylogeny packages. It can read and write trees in standard formats (PHYLIP, NEXUS, and CAIC). It allows drag-and-drop editing of branches, cut and paste of trees between windows (as NEXUS text), rerooting by clicking on branches, editing species labels, rotating branch order at polytomies, including and excluding species, and transforming branch lengths (including Sanderson's non-parametric rates smoothing) and many other interactive features. TreeEdit is available as an executable for Mac OS 8 or later at its web site at http://tree.bio.ed.ac.uk/software/treeedit/


[TreeThief icon]  Andrew Rambaut, of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) wrote TreeThief version 1.0, a program for inputting phylogenetic trees, with branch lengths, into the computer for use in other programs. The tree is input by clicking on each node in turn. A scanned image of a tree can be loaded first to act as a template. TreeThief is distributed from its web page at http://microbe.bio.indiana.edu:7131/soft/iubionew/molbio/evolution/phylo/TreeThief/ as a Mac OS executable. It requires a Macintosh running Mac OS 8 or later.


Thomas Laubach and Arndt von Haeseler of the Bioinformatics Institute of the University of Duesseldorf, Germany (thomas.laubach (at) uni-duesseldorf.de) and the Center of Integrative BioInformatics Vienna at the University of Vienna, Austria (arndt.von.haeseler (at) univie.ac.at) have produced TreeSnatcher Plus, a program to capture phylogenies from images. TreeSnatcher Plus is a GUI-driven Java program for the semi-automatic recognition of multifurcating phylogenetic trees in pixel images. The program accepts an image file as input and analyzes the topology and the metrics of a tree depicted, with user assistance. The analysis is carried out in a multiple-stage process using basic algorithms from the field of image analysis. It yields a tree in the standard Newick format that represents the tree structure, optionally including branch lengths. TreeSnatcher Plus can process trees with more than 100 leaves or more in a few seconds. It also allows the user to construct their own trees and to modify existing trees. An earlier version, Treesnatcher 1.0, is described in the paper: Laubach, T. and A. von Haeseler. 2007. TreeSnatcher: Coding trees from images. Bioinformatics 23: 3384-3385. It is available as Java executables, packaged for Windows, for Linux, and for Mac OS X. Its web site is available at two locations, at the University of Düsseldorf at http://www.cs.uni-duesseldorf.de/AG/BI/Software/treesnatcher/ and at the University of Vienna at http://www.cibiv.at/software/treesnatcher/. The same versions are available at both sites.


[TreeExplorer Icon] Koichiro Tamura of the Evolutionary Genetics Laboratory, Tokyo Metropolitan University, Tokyo, Japan (koichiro-tamura  (at) c.metro-u.ac.jp) has written TreeExplorer version 2.12, a program for plotting, rearranging, and editing trees, including a feature to compress a subtree to make essential features more visible. TreeExplorer is intended to work with MEGA but can also be used to read and edit or view trees in other standard formats. It is distributed as a Windows executable from its web site at http://evolgen.biol.metro-u.ac.jp/TE/TE_man.html where a manual for the program and a download link will be found.


[Mesquite icon] A (large) group headed by Katherine St. John at Lehman College, City University of New York (stjohn (at) lehman.cuny.edu), David Hillis and Tandy Warnow of the University of Texas, Austin (DHillis (at) mail.utexas.edu and tandy (at) cs.utexas.edu), and including numerous other people, has released Tree Set Viz, a program in Java for visualizing distances between sets of trees in a space. It computes distances between trees and uses these to infer points in a two-dimensional space that most closely approximate those distances. It is to be used in the Mesquite Java project for interacting with trees. Tree Set Viz is described in a paper: Hillis, D. M., T. A. Heath, and K. St. John. 2005. Analysis and visualization of tree space. Systematic Biology 54: 471-482. It is available from its web site at http://comet.lehman.cuny.edu/treeviz/software.html


[Treeme icon]  Dominik Hepperle of Sequentix, Dorfstrasse 20, D-18249 Klein Raden, Germany (info (at) do-mix.de) has released TreeMe, a program for visualisation and annotation of phylogenies. It can read and write trees, allows many kinds of annotation of nodes, rearrangement and editing of the tree, addition of multiple bootstrap values, collapsing of nodes, and more. It is available as a Windows executable from its web site in the company web sites at http://science.sequentix.de/software_treeme.php. It is purchased for 499 Euros for the first license, 199 Euros for each additional license, and 149 Euros for upgrade licenses.


[TreeGraph 2 icon]   Jörn Müller, of Pipinstraße 2, Bonn, Germany, Ben Stöver of the Nees-Institut für Biodiversität der Pflanzen, Rheinische Friedrich-Wilhelms-Universität Bonn, Germany (stoever (at) bioinfweb.info) and Kai Müller, of the group on Evolution and Biodiversity of Plants at Westfälische Wilhelms-Universität Münster, Germany (kaimueller (at) uni-muenster.de) have written TreeGraph 2 (version 2.0.45-197 beta), an editor for phylogenetic trees. It allows automatically combining information from different phylogenetic analyses and helps to identify and graphically present incongruencies. It features editing and formatting options, such as automatically setting line widths or colors according to the value of any of the unlimited number of variables that can be assigned to each node or branch. These node/branch data can be imported from spread sheets or other trees, be calculated from each other, be kept invisible or set visible and then be freely formatted. Whole clades can be copied from other files and be inserted, and can also be manually added. It can read NEXUS or Newick tree files, or its own TGF format, which uses a LaTeX-style format that is like those formats, but extended to indicate support values for nodes as well as annotations for groups of tips or clades. The program produces EPS, SVG, PDF or PNG graphics file formats. It is described in the papers:

TreeGraph 2 is Java code, available both as Java executables and as Java source code at its web page at http://treegraph.bioinfweb.info/. An earlier version, TreeGraph version 1.0 rc4, is available as source code for Unix or Linux, and as a Windows executable. It is available from its web site at http://www.math.uni-bonn.de/people/jmueller/extra/treegraph/.


   Adrian Canutescu and Roland Dunbrack of the Basic Science Division at the Fox Chase Cancer Center (Roland.Dunbrack (at) fccc.edu) have released ArboDraw version 1.0, a Windows program for creating, rendering, coloring, annotating, and saving images of phylogenetic trees. ArboDraw (from Latin, arbor, tree) is a Windows program for creating publication-quality images of phylogenetic trees. ArboDraw provides a combination of features:

  1. editing of the rendered tree, including independent horizontal and vertical scaling, full font control, and background and tree coloring;
  2. branch selection and coloring so that separate parts of the tree can be colored differently;
  3. annotation of sequences from the input FASTA header lines available on mouseover; labels and annotations can be edited as desired;
  4. control of saving the image in png format at desired resolution for publication;
  5. saving of the image in a novel XML format so that editing can be re-started from a previous editing session;
  6. running a fast and accurate sequence alignment program, MUSCLE, for generating trees; A MUSCLE executable is included with ArboDraw.
  7. alternatively allowing user input tree data in Newick format.
It is available as Windows executables which require the .Net 2.0 framework (freely downloadable from Microsoft). Arbodraw itself can be downloaded from its web site at http://dunbrack.fccc.edu/ArboDraw


François Chevenet and colleagues, of the Laboratoire de Génétique et Evolution des Maladies Infectieuses of the Institut de recherche pour le développement, UMR CNRS/IRD 2724, Montpellier, France (f.chevenet (at) ird.fr) have written TreeDyn version 198.3, modules for dynamic graphics and annotation of trees. TreeDyn can import trees, and place annotations on their nodes, branches, and clades. It can compare trees and find parts of trees which are similar to a highlighted region on a target tree. Operations in TreDyn can be scripted. It can handle very large trees and can collapse regions on those trees to concentrate on certain parts of the tree. TreDyn can handle multiple windows and multiple trees per window, can save and restore graphical analysis, can project texts or symbols facing leaf labels or linked to nodes. It includes a library of packages dedicated to specific research fields involving trees. It is described in the paper: Chevenet, F., C. Brun, A. L. Banuls, B. Jacq and R. Christen. 2006. TreeDyn: towards dynamic graphics and annotations for analyses of trees. BMC Bioinformatics 7: 439. It is available as source code in the graphics-oriented scripting language Tcl/Tk. There are separate distributions for Windows, for Linux, and for Mac OS X, for systems that already have Tcl/Tk installed. It can be downloaded from its web site at http://www.treedyn.org/


Robin Kramer, then of the Digital Biology Laboratory, Computer Science Department of the University of Missouri, Columbia, Missouri (whose email address was at that time: kramer (at) mizzou.edu) has produced DigTree (Digital Tree). DigTree is a tool that allows biologist to display large heirarchical trees in ways that preserve weights such as branch lengths. It can also display other sorts of trees. It is described in the paper: Kramer, R., V. Olman, Y. Xu, and D. Xu. 2006. DigitalTree: A tool for displaying biological data in tree structure. Lecture Notes in Computer Science vol. 3992, pp. 855-862. It is available as C++ source code and Windows executables. To compile it, one also needs the spirit++ framework in addition to a C++ compiler. TreeDyn can be downloaded from its web site at http://digbio.missouri.edu/~digtree/


Olivier Elemento, then of the LIRMM (Laboratoire d'Informatique de Robotique et de Microélectronique de Montpellier) at the Université de Monpellier II, Montpellier, France and now of the Institute for Computational Biomedicine at Weill Cornell Medical College, New York Citys (ole2001 (at) med.cornell.edu), has released DTdraw (Duplication Tree Draw), a program to draw trees of gene duplication histories. a command line tool to draw tandem duplication histories. Tandem duplication trees can be reconstructed with the DTscore program. DTdraw takes as input a (rooted) duplication history in Newick format, and an order file, i.e. a file which contains the number of leaves and the leaf labels of the tree, in correct order. At the moment, DTdraw outputs drawings in two graphical formats : PNG and Postscript. It is available as C source code and Linux executables. It can be downloaded from its web site at http://www.lirmm.fr/~elemento/DTdraw/


   Paul Michael Agapow, of the Department of Biology of Imperial College at Silwood Park, Ascot, Berks U.K. (agapow (at) agapow.net) has written TreeMaker, version 1.3, a program for interactive construction of taxonomies with species richness data. It allows the user to create a taxonomy and write out a tree that has species richness information on it. It is described in the paper: Crozier, R. H., L. J. Dunnett, P.-M. Agapow. 2005. Phylogenetic biodiversity assessment based on systematic nomenclature. Evolutionary Bioinformatics Online 1: 11-36. It is available as C source code, Windows executables and Mac OS X executables. It can be downloaded from its web site at http://www.agapow.net/software/treemaker. (TreeMaker is not related to this program of the same name)


Mike Sanderson of the Department of Ecology and Evolutionary Biology at the University of Arizona, Tucson, Arizona (sanderm (at) email.arizona.edu) has written Paloverde version 1.1, a program to display a phylogeny in one of several 3-dimensional formats. Paloverde is best used for trees of 100 to 2,500 tips. It reads the tree from a NEXUS tree file and can display it in a number of ways. These include a circular format, on a spiral, a cone, a hemisphere or in a three-dimensional treelike form. The user can rotate the object and zoom on it, and clades can be selected to be plotted. Subclades can also be collapsed to single tips. It is available as C source code and Powermac Mac OS X executables. It can also be compiled from the source code on Linux systems. It requires the free OpenGL graphics library. It can be downloaded from its web site at http://loco.biosci.arizona.edu/paloverde/paloverde.html


  Alessandro Zuccon and Dario Zuccon of the Department of Vertebrate Zoology at the Naturhistoriska riksmuseet, Stockholm, Sweden have produced MrEnt, version 2.2, a program to produce drawings of phylogenies. It is intended for drawing high-quality, printable phylogenetic trees. It is intended as a user-friendly program to recognize Nexus tree files and display them in the most commonly used tree formats. Trees can be displayed as rectangular cladograms, slanted cladograms or phylograms. The program allows full editing of taxon names, taxon comments, support values, and branches, addition of other graphic elements such as lines, shapes, text, brackets). Drawings can be exported in Portable document format (PDF), Windows enhanced metafile (EMF), Windows bitmap (BMP), or Portable Network Graphics (PNG) formats. It also allows multiple trees, layers and embedded graphics. It is available as a Windows executable. It also requires that the Microsoft .NET framework be installed. It can be downloaded from its web site at http://www.mrent.org"


[FigTree icon]  Andrew Rambaut of the Institute of Evolutionary Biology of the University of Edinburgh, Edinburgh, Scotland (a.rambaut (at) ed.ac.uk) has produced FigTree version 1.3.1, a program to graphically view phylogenies and prepare publication-ready figures from them. FigTree is designed as a graphical viewer of phylogenetic trees and as a program for producing publication-ready figures. Andrew notes that it was written for his own needs so may not be as polished and feature-complete as a commercial program. In particular it is designed to display summarized and annotated trees produced by BEAST. It is available as Java executables and Windows executables. It is available from its web site at http://tree.bio.ed.ac.uk/software/figtree/


[HyperTree icon]  Jonathan Bingham, then of Sugen, Inc. (more recently at Jivan Biologics: jbingham (at) jivanbio.com) wrote HyperTree version 12, a phylogenetic tree viewer, with a hyperbolic ('fish-eye') view and editing abilities that help in managing very large trees. HyperTree includes many features for navigating large trees, such as the ability to visualize large trees with hundreds of nodes or more, to rotate and drag the display in cartesian space, to search and select nodes, to copy clusters for pasting into other programs, to color-code branches, to zoom in and out, and to label branches (eg, common family members). It can view phylogenetic trees and other hierarchical clusterings, such as gene expression profile clusters. The program subsequently came to be owned by Pfizer, Inc., which has given permission to the Salk Institute for its distribution at the kinase.com web site. It is described in the paper: Bingham, J. and S. Sudarsanam. 2000. Visualizing large hierarchical clusters in hyperbolic space Bioinformatics 16(7): 660-661. It is available as Java executables. It can be downloaded from its web site at http://kinase.com/tools/HyperTree.html


[Dendroscope icon]  Daniel Huson, with contributions from Tobias Dezulian, Markus Franz, Christian Rausch, Daniel Richter and Regula Rupp of the Center for Bioinformatics of the Tuebingen University, Tuebingen, Germany (huson (at) informatik.uni-tuebingen.de) has released Dendroscope 2, an interactive program for the visualization and navigation of phylogenetic trees. Dendroscope is a new program for the interactive visualization and navigation of phylogenetic trees. The program provides eight standard tree visualizations and is optimized to handle trees containing hundreds of thousands of taxa. The program allows trees to be edited as well as for graphics to be exported. Version 2 adds the capability of displaying networks as well as trees. To support the inspection of large trees, Dendroscope offers a magnification tool. It is written in Java and and installers are provided for Linux/Unix, MacOS X and Windows. It is described in the paper: Huson, D. H., T. Dezulian, M. Franz, C. Rausch, D. C. Richter and R. Rupp. 2007. Dendroscope: an interactive viewer for large phylogenetic trees, submitted for publication. Dendroscope can be downloaded from its web site at http://www-ab.informatik.uni-tuebingen.de/software/dendroscope for free, although the authors require that you obtain a free license key online and cite their paper in any publication using the program.


[PhyloWidget icon] Gregory Jordan of the Goldman Group of the European Bioinformatics Institute (phylowidget (at) gmail.com) or greg (at) ebi.ac.uk) has written PhyloWidget version 1.0, a web-friendly tool for creating, editing, and visualizing trees. PhyloWidget is a phylogenetic tree visualization program that "plays nice" with the Internet. It runs directly from the browser window (as a Java applet), and trees can be loaded in manually from the user's hard drive, or alternatively through a URL-based API. They can be displayed and edited, and publication-quality figures produced. PhyloWidget is aimed at (1) users who want a simple, easy-to-use tree visualization tool without having to download software, and (2) phylogenetic tree databases who wish to use the URL API to let PhyloWidget visualize the database's trees. It is described in the paper: Jordan, G. E. and W. P. Piel. 2008. PhyloWidget: Web-based visualizations for the tree of life. Bioinformatics 24: 1641-1642. It is available as Java source code and Java executables, and a browser-based Java applet. It can be downloaded from its web site at http://www.phylowidget.org/


John Brzustowski of the Department of Biological Sciences of the University of Alberta, Edmonton, Canada (jbrzusto (at) ualberta.ca) has written TreeToy , a program to view a tree and select subtrees from it. TreeToy reads a tree from a file and displays it. The user can reroot the tree or select subtrees to write out. It can also be zoomed and written out in a PICT2 format for printing. It is available as Java source code and Java executables. It can be downloaded from its web site at http://www2.biology.ualberta.ca/jbrzusto/TreeToy.php


[Phyutility icon]   Stephen A. Smith and Casey W. Dunn, of the Department of Ecology and Evolutionary Biology of Brown University, Providence, Rhode Island (stephen_a_smith (at) brown.edu) have produced Phyutility version 2.2, a utility to modify trees and to perform certain data set modifications as well. Phyutility's tree operations include rerooting, pruning, type conversion, consensus trees, measures of leaf stability, of the varying attachment locations of lineages, and measures of tree support. It can also do some data operations such as concatenating alignments, parsing Genbank files, trimming alignments, and searching for and fetching sequences from the Genbank database. It is described in the paper: Smith, S. A. and C. W. Dunn. 2008. Phyutility: a phyloinformatics tool for trees, alignments, and molecular data. Bioinformatics 24: 715-716. It is available as Java source code and Java executables. It needs two additional Java libraries downloadable from elsewhere. It can be downloaded from its web site at http://code.google.com/p/phyutility/


Catherine Anderson of the Department of Computer Science of the University of Nebraska, Lincoln (cateander27 (at) gmail.com) has released SuiteMSA (A suite of tools for visual comparision of Multiple Sequence Alignments), version 1.3.14, Contains tools that allow the visual and quantitative comparison between multiple sequence alignments using sequence sets as well as structural information. . This package contains tools that allow the visual and quantitative comparison between MSA using sequence sets as well as structural information. Various color scheme are available including color schemes based on structural information. The Pixel Plot tool allows the display of multiple large alignment in a pixelated display that gives the overall shape of the alignments as well as column wise comparison. Included in this package is a sequence simulator, iSGv2. It can also act as a GUI and call the alignment programs ClustalW and MUSCLE, and it also has as a phylogeny display. It is described in the paper: Anderson, C. L., C. L. Strope and E. N. Moriyama. 2011. SuiteMSA: Visual tools for multiple sequence alignment comparison and molecular sequence simulation. BMC Bioinformatics 12: 184. It is available as Perl script, Java executables and Intel Mac OS X executables. It can be downloaded from its web site at http://bioinfolab.unl.edu/~canderson/SuiteMSA/


David Kidd and Xianhau Liu of the National Evolutionary Synthesis Center at Duke University, Durham, North Carolina (dk (at) nescent.org) have released GeoPhyloBuilder version 1.0.2, an extension for the widely-used geographic information system program ArcGIS that creates geophylogenies. Geophylogenies are spatial networks in which network topology is defined by a phylogenetic model and the geographical position of nodes and branch paths are derived from a biogeographical model. Tree depths are assigned to the z-dimension of the spatial network facilitating rapid 3D visualization in GIS or other software. Geophylogenies can be output in ArcGeodatabase, shapefile and KML (Google Earth) format. A copy of ArcGIS is required. GeoPhyloBuilder is described in the paper: Kidd, D. M. and X. Liu. 2007. geophylobuilder 1.0: an ArcGIS extension for creating 'geophylogenies'. Molecular Ecology Notes 8 (1): 88-91. It is available as Windows executables and source code for the Microsoft C++ .NET compiler. It can be downloaded from its web site at http://www.nescent.org/informatics/software.php


[GenGIS icon]   Donovan Parks and Robert Beiko of the Faculty of Computer Science at Dalhousie University, Halifax, Nova Scotia, Canada (beiko (at) cs.dal.ca) have released GenGIS version 1.08, a Geospatial Information System for Genetic Data. GenGIS allows the integration of digital map data alongside information about sequences and sample locations. A wide range of data visualizations are supported, and phylogenetic trees or cluster relationships can be drawn in two or three dimensions. A Python interface provides extensibility and camera controls, while the R statistical package allows exposed data to be subjected to statistical analysis. It is described in the paper: Parks, D. H., M. Porter, S. Churcher, S. Wang, C. Blouin, J. Whalley, S. Brooks, and R. G. Beiko. 2009. GenGIS: A geospatial information system for genetic data. Genome Research 19: 1896-1904. It is available as C++ source code, Windows executables and Mac OS X universal executables. It can be downloaded from its web site at http://kiwi.cs.dal.ca/GenGIS/Main_Page


Richard H. Ree and Stephen A. Smith of the Department of Botany of the Field Museum of Natural History (rree (at) fieldmuseum.org) has released Lagrange (Likelihood analysis of geographic range evolution), version 2.0.1, likelihood models for geographic range evolution on phylogenetic trees, with methods for inferring rates of dispersal and local extinction and ancestral ranges. Lagrange (likelihood analysis of geographic range evolution) is a freely available Python package, co-authored by Ree and Smith, implementing dispersal-extinction-cladogenesis models of geographic range evolution, methods described in papers by Ree et al. (Evolution, 2005) and Ree and Smith, 2008. It can be used to infer geographic ancestry (historical ranges) and estimate rates of dispersal and local extinction on phylogenetic trees that have species range data arrayed at their tips. A web server that is an analysis configuration tool that prepares files to be run on your copy of Lagtange is available at: http://www.reelab.net/lagrange. Lagrange is described in the paper: Ree, R. H. and S. A. Smith. 2008. Maximum likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Systematic Biology 57(1): 4-14. It is available as C++ source code and Python script. It can be downloaded from its web site at http://code.google.com/p/lagrange/


[Phylocom icon]  Campbell Webb, David Ackerley, and Steven Kembel of the Arnold Arboretum of Harvard University, of the University of California, Berkeley, and the Center for Ecology and Evolutionary Biology of the University of Oregon (cwebb (at) oeb.harvard.edu) have written Phylocom (software for the analysis of PHYLOgenetic COMmunity structure and character evolution) version 4.1, a program to measure phylogenetic community structure and carry out comparative methods analysys. It can measure community phylogenetic structure (Webb's mean phylogenetic distance, mean nearest phylogenetic neighbour distance, NRI and NTI, Faith's phylogenetic diversity, phylogenetic distances among samples and taxa) and compare observed patterns to those expected under various null models of community assembly and phylogeny structure. It cna also carry out comparative methods for the analysis of character evolution and speciation (independent contrasts, lineages through time, and trait evolution under different evolutionary models). In addition it can manipulate phylogenetic trees and ecological community data (converting Newick to Nexus format, pruning and merging phylogenetic trees, and doing data randomization using one of several null models). It can turn species lists into phylogenies with branch lengths based on fossil age estimates. It is available as C++ source code, Windows executables and Mac OS X universal executables. It can be downloaded from its web site at http://www.phylodiversity.net/phylocom/


Gabriel Valente and Rod Page of the Department of Software of the Technical University of Catalonia, Barcelona, Spain and the Division of Environmental and Evolutionary Biology of the Institute of Biomedical and Life Sciences at the University of Glasgow, Scotland (valiente (at) lsi.upc.edu) have produced Forest version 0.1, programs for producing or using edit scripts to convert one tree to another in the GML graph representation language. GML, the Graph Modelling Language (not to be confused with the Geographic Markup Language GML) is a file format for graphs. Forest contains programs to take two trees written in GML format, and produce an edit script that can convert one into the other. It also contains a program to apply the script to a tree. It is described in the paper: Page, R. D. M. and G. Valiente. 2005. An edit script for taxonomic classifications. BMC Bioinformatics 6: 208. It is available as C++ source code. i To compile this the user also needs to have the Graph Template Library. Forest can be downloaded from its web site at http://darwin.zoology.gla.ac.uk/~rpage/forest/


  Steven Brewer (sbrewer  (at) bio.umass.edu) of the Biology Department, University of Massachusetts, Amherst, Massachusetts, and Robert Hafner, formerly of Western Michigan University have developed Phylogenetic Investigator version 2.01, a teaching program that allows students to connect together organisms from a data set provided to them, to make phylogenies and examine them. The program is a Mac OS executable. Phylogenetic Investigator and its manual are downloadable here at http://bioquest.org/BQLibrary/library_details.php?product_id=22909. It is described in two papers:

Phylogenetic Investigator appears in BioQUEST Library VI) from the BioQUEST consortium, (whose web page is at http://bioquest.org), a nonprofit publisher of interesting biological teaching software. Although in the past the BioQUEST Library modules were only available on a CDROM sold by them, they are now freely downloadable.


David Joyce, of the Department of Mathematics and Computer Science, Clark University, Worcester, Massachusetts (djoyce  (at) clarku.edu) makes available the Phylap program, version 0.2, a Java applet that teaches about inferring phylogenies from molecular sequences using some clustering algorithms. The program allows the user to choose some parameters including the number of species, the rate of evolution and how many states there are at sites in the molecule. The program then chooses a random tree and evolves sequences on it, then computes distances between the sequences and infers the tree by one of several clustering methods. The user can see the results at each stage. The java applet may be downloaded from its web site at http://babbage.clarku.edu/~djoyce/java/Phyltree/cover.html, and if your browser can run Java programs it can of course also run the applet when browsing that page.


  I (Joe Felsenstein, Department of Genome Sciences and Department of Biology, University of Washington, Seattle, joe  (at) gs.washington.edu) have written a program, Dnatree version 1.3, for teaching about molecular phylogenies. The program uses a crude character-mode (not GUI) user interface. It is based on several programs from PHYLIP. The user chooses some rate parameters and the number of species. The program then simulates a random tree using a birth process and displays it. The user can then have the program simulate evolution of a stretch of DNA sequence along this tree. Then the program displays a different random tree, showing the number of changes inferred along that tree by a parsimony criterion. It allows the user to manually rearrange the tree to try to minimize the number of changes. Dnatree is available as C source code and as executables for Linux, Windows, and Mac OS X from its web site at http://evolution.gs.washington.edu/dnatree/dnatree.html Some executables of an earlier version, 1.1, are also available there for Mac OS, 68k Macintoshes, SunOS, Compaq/Digital Alpha, and IBM AIX.


Dave Dobson of the Geology Department of Guilford College, Greensboro, North Carolina (ddobson (at) guilford.edu) has written SimpleClade, a program that allows simple cladistic analysis with a graphical user interface. SimpleClade takes a data file in a common spreadsheet format, with values in the columns which are treated as identifiers of discrete states. It displays an arbitrary tree and then allows you to manually rearrange it interactively. The parsimony score is displayed as you do so. Outgroups can be chosen and the directions of the state changes are displayed as well as the tree. The program was written primarily as a teaching tool. It is available as a Windows executable. It can be downloaded from its web site at http://www2.guilford.edu/geology/simpleclade/


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