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Hipp Marcial Escudero ABSTRACT Summary: MaATiICCE is a new software package written in the R language that implements an information-theoretic approach to estimating the probability of continuous character transitions on phylogenetic trees. It relies on and provides helper functions for the automates batch-analysis of alternative models of character change utilizing the OUCH (R) R package; allows users to account for estimates the relative support for character changes having occurred at numerous nodes on a phylogeny while integrating over uncertainty in the phylogenyetic and the model of character change uncertainty; and provides discrete-time simulation functions for visualizing analysis results as well as a priori models. Availability: MaATiICCE is open source and written entirely in R language. Source and compiled binaries are freely available through RForge ( HYPERLINK "http://r-forge.r-project.org/" http://r-forge.rproject.org/ ). Contact: HYPERLINK "mailto:ahipp@mortonarb.org" ahipp@mortonarb.org INTRODUCTION Inferring the evolutionary history of species traits is a basic fundamental problem in evolutionary biology and an issue of practical concern for ecologists, developmental biologists, and others testing biological hypotheses using phylogenetic data. Methods of analyzing categorical data in a phylogenetic context have addressed the problem of localizing character transitions to branches {Huelsenbeck, 2003 #533;Ree, 2005 #1286;Nielsen, 2002 #535}. Transitions New methods utilizing phylogenetic generalized least squares model shifts in inthe dynamics of continuous character evolutions {Revell, 2009 #1444;O'Meara, 2006 #813;Scales, 2009 #1469;Butler, 2004 #598}. XXX. have received less focus, despite the fact that among-clade differences in character distributions often reflect a biologically interesting process (shifts in environment, changes in evolutionary dynamics or population-level processes, etc.). Alternatives to the constant variance model (Brownian Motion model) of continuous character evolution have until recently focused primarily on branch-length transformations that model how different evolutionary scenarios affect phylogenetic structure and the mode and tempo of evolution {Pagel, 1999 #432;Blomberg, 2003 #1312;O'Meara, 2006 #813}. These methods, however, are not generally useful for identifying (1) whether an apparent change in the distribution of a character on a tree is significant enough to warrant further investigation, (2) whether the particular distribution tells us something about the evolution of this trait, and (3) where on the phylogeny this change in distribution took place. Evaluating alternative Ornstein-Uhlenbeck (O-U) models in a phylogenetic context {Butler, 2004 #598;Martins, 1997 #421;Hansen, 2008 #1277} may be used to address such questions about evolutionary transitions in continuous characters. The use of O-U models typically focuses on testing adaptive scenarios {Bergmann, 2009 #1319;Smith, 2008 #1091;Verdu, 2006 #787} or phylogenetic constraints {Diniz-Filho, 2001 #987;Blomberg, 2003 #1312}. However, two recent studies {Hipp, 2007 #1051;Lumbsch, 2008 #1282} have utilized O-U models to estimate the probability of continuous character transitions on phylogenetic trees, modeled as shifts in character equilibrium or optimum. The approach taken entailed (1) identifying n nodes at which a change may have occurred, (2) evaluating support for all 2n models that allow all permutations of change at those nodes, and (3) estimating the support for a transition in character state at each node studied as the cumulative information criterion weight for all models entailing a shift at that node {Burnham, 2002 #791}. The software described in this application note—MmaatTicceCCE, a package written in the R statistical language {R-Development-Core-Team, 2004 #588}—extends this method by allowing for phylogenetic uncertainty and for limits on the maximum number of nodes at which transitions are allowed to occur; implementing a set of discrete-time simulations for visualization of character evolution under different models; providing an easy interface by which to test numerous whole-tree and partitioned-tree models for a particular phylogenetic branch; and allowing for flexibility in defining nodes for analysis. The method is similar to a stepwise AIC approach {Yamashita, 2007 #1532;Alfaro, 2009 #1509}. However, by taking a model-averaging approach rather than a model-selection approach, it is not susceptible to the significance threshold effects that introduce uncertainty into model-selection. IMPLEMENTATION The MmatTicceCCE package (http://r-forge.rproject.org/projects/mattice/) provides functions for generating the potentially large number of model specifications required to test character transition hypotheses; performing and summarizing analyses over a single tree or a set of trees, allowing analyses to be performed even for nodes that are not found in all trees being studied; simulating and visualizing character evolution under a user-specified model or using model-averaged parameters from a mMatTicceCCE analysis; and evaluating the relative support for several different character-transition models on a single node. The approach presupposes that (1) flat priors on all models are reasonable; (2) all nodes of interest have been specified; (3) XXX The likelihood calculations are all performed in ouch {Butler, 2004 #598;King, 2009 #1313}, but addressing these particular questions entails summarizing analyses over a large number of specified models and extracting tree data to analyze over phylogenetic uncertainty, functions that would be superfluous in ouch. AVAILABILITY For more details see the vignette: vignette(‘maticce’) Installation: install.packages("maticce", repos="http://R-Forge.Rproject.org") Q R S T V W Z ‘ ’ “ ³ ´ ½ Ä Å Æ Ç È É Ê Ë Ì õ ù û ôìÝÅìÝÅìôìÀ´¥– ‹ƒ~Àr]Àr]ÀQÀ • H h/ ÖF h:SJ ] • ( • h´0½ h;?2 ] •cH dh dh dh ÓF • H h ÓF h;?2 ] • há@X ] • há@X 5 •\ •] • há@X há@X 5 •\ •] • h;?2 há@X 5 •\ •] •mH U sH h;?2 h´0½ 5 •\ •] •mH sH h;?2 h´0½ ] •mH sH h´0½ ] •. • h´0½ h;?2 5 •\ •] •cH  ÓF h;?2 5 •\ •] • h´0½ 5 •\ •] • h´0½ Q ’ “ ¢ ³ ´ ½ } Z dh dh dh ÓF h´0½ 5 •\ •] • • H 8 h P £ ¤ ± ¦ § ý ¶ ¬ - ý ý èý ý ö- . ê •ý ý ý ø ê ê ý ý ý ê ÷ý ý ý ê é- ê ø ý ï ê ê gd€ ø c „Ð`„Ðgd¯+° û gd w - 9 Ž ’ ™ œ  ' C M f v w x À Ó ôïçïçïÒ½Ò¨œï•uï`THTHTH< • H h”Ú؆hº9ç ] • • H h“Ú؆hº9ç ] • • H h‘Ú؆hº9ç ] • ( hš3 h:SJ ] •cH dh dh dh0 ÖF 5 • • H hÄ ÓF h;?2 h:SJ ] •cH dh dh dh/ ÖF • H h/ ÖF h:SJ ] • • H hÚ؆hº9ç ] • ( • h´0½ hº9ç ] •cH dh dh dhÚ؆ ( • hõw • h:SJ ] •cH dh dh dh/ ÖF ( • h´0½ h:SJ ] •cH dh dh dh/ ÖF h´0½ h´0½ ] • h´0½ ] • • H hà ÓF h;?2 ] • Ó Û Ü à å é î D | } ‹ Œ • Ž • • ‘ ½ Æ 5 7 8 9 V W Z b c d Œ úîÙúîúîÙúÔúÏǶ¡Â¶¡Â†Â|ÂtÂd|Z|ÂÇÂ| h S± há@X 0J ] •- • j h S± há@X U ] • há@X há@X ] • j há@X U ] • 5 • • H hÄ ÓF h; ?2 h:SJ ] •cH dh dh dh0 ÖF( • há@X h;?2 ] •cH dh dh dhÄ ÓF • H hÄ ÓF h;?2 ] • há@X ] • há@X 5 •\ •] • h¥ ¥ ] • h´0½ ] •( hš3 hº9ç ] •cH dh dh dh”Ú؆ • H h”Ú؆h º9ç ] • hš3 ] • #Œ • Ž ¡ ¢ ¤ ° ± ë ñ ü ý 5 6 q s t • ¼ Â Ì Î Ý ðæÜæ×Ï×ǯ£”ÇŠÇu]QEQ]9H h1 ÖF h:SJ ] • • H h— Ú؆h|Eä ] • • H h2 ÖF h:SJ ] • . • hõw• hõw• h:SJ ] •cH dh d h dh2 ÖF ( • hõw• h:SJ ] •cH dh dh dh2 ÖF hö•~ ] • hõw• ] • • H h–Ú؆hõw• h|Eä ] • • H h– Ú؆h|Eä ] • . • hõw• hõw• h|Eä ] •cH dh dh dh– Ú؆ hõw• hõw• ] • há@X 5 •\ •] • há@X ] • h S± há@X 0J ] • j há@X U ] • - • j h S± há@X U ] • Ý î ò ü ý þ $ K P Q j • ˜ ¦ ´ å ë ì B D p ˆ ª ‹ Œ Ÿ à b c À Å 3 9 @ ÷òæÑòŹŹ-¹ò¥ ò¥ ˜ ˜ ˜ ˜¥“‡‚“}“‚u‚u‚u‚“m“m hõw• h€c ] • h qÊ h w ] • hB 6 ] • h w ] • • H hÆ ÓF h;?2 ] • h€c ] • h qÊ h:y¬ ] • h:y¬ ] • h qÊ h qÊ ] • • H h3 ÖF h&+ ] • • H h3 ÖF h:SJ ] • • H h2 ÖF h:SJ ] • ( • h qÊ h:SJ ] •cH dh dh dh1 ÖF • H h1 ÖF h:SJ ] • h qÊ ] • hõw• hõw• ] • *@ Y “ ñ ò È ª « µ Ã Ô Õ ! 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