paleotree: an R package for paleontological and phylogenetic analyses of evolution
|
|
- Johnathan Stafford
- 5 years ago
- Views:
Transcription
1 Methods in Ecology and Evolution 2012, 3, doi: /j X x APPLICATION paleotree: an R package for paleontological and phylogenetic analyses of evolution David W. Bapst* Department of Geophysical Sciences, University of Chicago, 5734 South Ellis, Chicago, IL 60637, USA Summary 1. paleotree is a library of functions for the R statistical computing environment dedicated to analyses that combine paleontological and phylogenetic data sets, particularly the time-scaling of phylogenetic trees, which include extinct fossil lineages. 2. The functions included in this library focus on simulating paleontological data sets, measuring sampling rates, time-scaling cladograms of fossil taxa and plotting historical diversity curves. 3. I describe the capabilities and analytical basis of the functions in paleotree by presenting two examples. The first example showcases the simulation capabilities and plotting the output as diversity curves. The second example demonstrates time-scaling a cladogram of fossil taxa and estimating sampling rates and completeness from temporal ranges. Key-words: birth-death models, comparative methods, diversification, macroevolution, paleobiology Introduction Increasingly, paleobiology and evolutionary biology are finding common ground in the shared realm of phylogenetics. Paleobiologists are increasingly applying phylogeny-based approaches, developed by evolutionary biologists, to study evolution in the fossil record (Fusco et al. 2012; Lloyd, Wang & Brusatte 2012). The last few years have seen a rise in the use of the free and platform-independent R coding environment (R Development Core Team, 2011) among members of the applied phylogenetics community. This movement has produced a wealth of R libraries dedicated to the analysis of phylogenies and evolutionary patterns (e.g. Paradis, Claude & Strimmer 2004; Harmon et al. 2008; Schliep 2011; Revell 2012). However, only a few libraries deal explicitly with issues encountered in analyses of paleontological phylogenies, such as paleophylo which focuses on depicting relationships among fossil taxa (Ezard & Purvis 2009). In this article, I present the R library paleotree, which implements methods for the analysis of paleontological data and the evaluation of models, with a particular focus on phylogenetic approaches. paleotree is designed to concentrate on four types of analyses: (a) simulating diversification and sampling in the fossil record, (b) calculating diversity changes across time from different data types, such as taxonomic ranges and phylogenies, (c) estimating sampling parameters from observed temporal ranges and (d) time-scaling cladograms of extinct taxa. In addition, paleotree offers a wide range of additional utilities *Correspondence author. dwbapst@uchicago.edu related to these areas, such as converting simulated fossil recordsintostandardphylogeneticdatastructures.paleotree can be installed from any of the mirrors for the Comprehensive R Archive Network, the standard repository for publicly-released R libraries. Many of the functions in paleotree depend on the functions and standards set by the free library ape (Paradis, Claude & Strimmer 2004). The S3 object class phylo established by ape gives programmers a structure, which can be used as a common currency among other phylogenetics packages in R, including the option to add modules to the phylo data structure (e.g. Revell 2012). The various time-scaling functions for paleontological phylogenies in paleotree require objects of class phylo as input and also output phylogenies in this format, as do the simulation-conversion functions taxa2phylo and taxa2cladogram. Functions in paleotree that produce time-scaled phylo objects will alter the standard format of these objects at output by adding a new element, $root.- time. This value informs other functions in paleotree at what time before present the root divergence occurred, as most functions in ape and other packages assume that the tip furthest from the root on a tree with edge lengths is at the modern time (zero time-units before present). This is useful, as many paleontological trees will have their tips located far from the present, such as a phylogeny of non-avian dinosaurs or trilobites. In addition to depending on ape, paleotree also imports functions from the publicly available libraries geiger (Harmon et al. 2008) and phangorn (Schliep 2011). I describe here a multi-part example that demonstrates several capabilities of the paleotree library. First, I obtain Ó 2012 The Authors. Methods in Ecology and Evolution Ó 2012 British Ecological Society
2 804 D. W. Bapst a simulated data set of taxonomic ranges from a birth death model, model sampling in these lineages and convert the simulation into phylogenetic data sets. I also plot the diversity curves calculated from these different versions of the same simulation. In the second part of the example, using the simulated data set from the first part, I demonstrate the functions in paleotree for time-scaling cladograms of fossil taxa using their ranges. Finally, I discuss using the sampling rate conditioned ( SRC ) time-scaling methods, including an example of estimating the prerequisite instantaneous sampling rate from taxon ranges. Example Part 1: simulating fossil data and plotting diversity curves The main simulation function in paleotree is simfossil- Taxa, which simulates diversification under a birth death model (Kendall 1948; Nee 2006). There are a number of birth death simulators available via R, such as the function rlineage in ape. simfossiltaxa differs by enmeshing the birth death process with models of morphological differentiation across lineages. This is important for simulating paleontological data sets, as the interpretation of evolutionary patterns in the fossil record is dependent on the recognition and identification of morphologically-defined taxa. By manipulating the function s arguments, several different patterns of morphological differentiation can be simulated (Wagner 1995; Wagner & Erwin 1995; Foote 1996). Differentiation can occur in both daughter lineages of branching events ( bifurcating cladogenesis ), in only a single daughter lineage ( budding cladogenesis ) or in neither descendant lineages ( cryptic cladogenesis ). Shifts in morphology can also be modeled along branches, independent of branching events ( anagenesis ). simfossiltaxa can simulate any mixture of these processes by treating these as discrete events leading to the recognition of new taxa, instead of directly simulating morphological change. By default, clades are simulated with no anagenesis and pure budding cladogenesis at each branching event. The selected model of morphological differentiation can greatly impact some analyses while having little effect on others, such as estimates of diversity. The output of simfossiltaxa is a matrix describing the ancestor descendant relationships and temporal ranges of morphologically distinct taxonomic units. In addition to the options available for simulating morphological divergence, the acceptance of simulation runs in simfossiltaxa can be conditioned on a number of various minimum and maximum criteria. Simulation runs are accepted for output if and when they meet all of these criteria. The code below simulates a clade using typical birth death parameters (speciation rate = extinction rate = 0Æ1 per lineage time-units), conditioned so that the resulting output has between 120 and 150 morphologically distinguishable taxa over its evolutionary history and no living taxa at the end of the simulation. The diversity curve of this data is plotted, depicting the true history of diversity changes for our simulated clade. library(paleotree) # load the library set.seed(1) # set random seed taxa <- simfossiltaxa(p=0æ1,q=0æ1,nruns=1, mintaxa=120, maxtaxa=150, maxtime=1000,maxextant=0) # plot true diversity curve as Fig. 1A layout(1:3);par(mar=c(4æ5,4,4,1)) taxadiv <- taxicdivcont(taxa) # saves div curve data as taxadiv title("true Diversity Curve") The function sampleranges can be used to simulate incomplete sampling across a set of lineages output by sim- FossilTaxa. This function stochastically places sampling events along the original ranges and outputs the temporal ranges that would be observed given those samples. By default, sampleranges assumes a model where sampling events are independently distributed across branches and throughout time as a Poisson process (Foote & Raup 1996; Foote 1997), with some instantaneous rate. By altering the function s parameters, sampling can be simulated under more complex, time-varying models (Liow, Quental & Marshall, 2010). In the example below, the instantaneous sampling rate is set to 0Æ2 per lineage time-units. The function bintimedata takes sampled continuous-time ranges as input and returns these same ranges on a discrete interval time-scale, here with intervals five time-units long. So that all three diversity curves are plotted against the same horizontal time-scale, we will directly input the interval times saved from calculating the true diversity curve above. # sample ranges rangescont <- sampleranges(taxa,r=0æ2) # plot figure 1B taxicdivcont(rangescont, int.times=taxadiv[,1:2]) title(c("sampled Diversity Curve", "(Continuous Time)")) # bin ranges into discrete intervals rangesdisc <- bintimedata(rangescont, int.length=5) # plot figure 1C taxicdivdisc(rangesdisc, int.times=taxadiv[,1:2]) title(c("sampled Diversity Curve", "(Discrete Time)")) The resulting figure reveals that although the diversity histories estimated from the sampled ranges have a similar pattern of increases and decreases as the original data set, the undersampled simulation has less diversity observed per time-unit (Fig. 1). The simulated fossil record can also be converted into a phylogeny using two different functions, depending on what type of data is desired. taxa2phylo constructs the time-scaled tree that perfectly describes the set of relationships for particular points in time within the simulated taxon ranges. Taxonomic identity of branches is lost; only the historical patterns of branching matter. Ancestral taxa with multiple descendants
3 Analyses of Paleo-Trees in R True diversity curve Sampled diversity curve (continuous time) Sampled diversity curve (discrete time) distic analysis with a very large number of informative characters, only capturing the true resolvable relationships among a simulated set of taxa. Output from taxa2cladogram is generally poorly resolved, as the sampling of ancestral taxa and static taxa with multiple descendants will produce non-nesting relationships that are not resolvable using typical morphological phylogenetics and thus cannot be portrayed on a cladogram (Smith 1994; Wagner & Erwin 1995). cladogram <- taxa2cladogram(taxa,plot=t) The diversification histories of fossil taxa in paleotree can be further investigated by examining the phylogenetic structure among lineages extant at specific dates in the history of a clade. This is possible with the function timeslicetree that removes the portions of a phylogeny that passes some specified point in time. For this example, the tree is sliced at 800 time-units before present, during a diversity peak in the simulated clade. Terminal branches that have gone extinct by that date can be dropped using the argument drop.extinct, allowing us to simulate the type of data generally available to biologists. timeslicetree outputs an ultrametric phylogeny, which can be visualised as a lineage-through-time plot with phylodiv Fig. 1. Three plots of the changes in taxonomic diversity for a simulated clade. The first is the true record of the evolutionary history of these taxa, and the second is the result of a simulation of sampling on the true ranges, in continuous time. The third plot is a conversion of the sampled continuous-time ranges onto a time-scale of discrete intervals, each five time-units long. are chopped into multiple segments to become multiple branches within the output phylogeny. Tip taxa represent the position of the per-taxon instantaneous observations; by default, these are the last occurrences of each taxon, but this can be changed with the argument obs_time. In general, users should use taxa2phylo in simulation studies, such as when simulating trait data with functions that require phylo objects. tree <- taxa2phylo(taxa) # get diversity curve phylodiv(tree,int.times=taxadiv [,1:2]) Alternatively, taxa2cladogram produces an unscaled cladogram that contains the set of nesting relationships that are resolvable with morphological data among the input taxa, given the pattern of morphological shifts and ancestor-descendant relationships in the input. The result emulates an ideal cla- Lineage richness (log scale) Fig. 2. The phylogeny at top depicts the phylogenetic relationships along taxa present at 800 time-units before present in the simulated clade, with extinct lineages removed, obtained using the function timeslicetree. The figure below it is a plot of the increase in lineage diversity reconstructed from the phylogeny above, with richness plotted on a log-scale (numbers on the axis are actual richness, not log-richness).
4 806 D. W. Bapst tree800 <- timeslicetree(tree, slicetime=800,drop.extinct=t) # LTT plot (Figure 2) phylodiv(tree800,plotlogrich=t) Example Part 2: estimating sampling rates and time-scaling paleontological phylogenies Often, workers who wish to use phylogenetic approaches for studying evolution in the fossil record begin with an unscaled cladogram and a set of taxon ranges. To further complicate matters, fossil taxa are generally known only as first and last appearances in discrete temporal intervals, adding temporal uncertainty to when they occurred, and the cladograms may be partially unresolved. In paleotree, the function time- PaleoPhy time-scales trees, using any of several typically applied time-scaling methods (e.g. Hunt & Carrano 2010; Hopkins 2011; Lloyd, Wang & Brusatte 2012). The default time-scaling method, referred to as the basic method in the documentation, sets node ages equal to the first appearance time of their earliest descendant taxon (Smith 1994). timepaleophy is specialised for data sets where taxon occurrences are known in continuous time, but paleotree also offers bin_timepaleophy,a wrapperfor timepaleo- Phy that accepts taxon occurrences in discrete time intervals. When bin_timepaleophy is used, per-taxon first and last occurrence dates are pulled from a uniform distribution, bounded by the upper and lower bounds of the intervals from which those occurrences are listed. This continuous-time data set, stochastically built from the discrete-time input, is then used as input for timepaleophy. If a pair of dates would imply a negative duration for a taxon, they are discarded and drawn again. This is similar to methods employed by Lloyd, Wang & Brusatte (2012). To demonstrate the time-scaling capabilities of paleotree, I will give an example of time-scaling the ideal cladogram simulated above with the discrete-time ranges, as the majority of paleontologists using this package will only have occurrences in discrete intervals. timetree <- bin_timepaleophy(cladogram, rangesdisc) # plot the tree plot(ladderize(timetree),show.tip.label=f) In the example above, bin_timepaleophy output a timescaled phylogeny where the tips are placed at the first appearance dates of the included taxa. In other words, the observed terminal portions of those taxonomic ranges were not added onto the output tree. This default setting can be changed using the argument add.term.typically,terminal branches are not added in analyses of characters considered static over taxon ranges, but should be included in analyses of diversification. timetree <- bin_timepaleophy(cladogram, rangesdisc,add.term=t) #plot the tree plot(ladderize(timetree),show.tip.label=f) By default, bin_timepaleophy outputs only a single time-scaled tree. As the occurrence dates used are stochastically drawn from uniform distributions, this function automatically returns a warning that strong interpretations should not be made on a single output tree because of this stochastic element. Analyses based on a single tree may produce misleading results. Users should instead consider a sample of trees with different dates stochastically chosen, by increasing the ntrees argument to a suitably large sample size. When ntrees is increased, the output is of the ape class multiphylo (Paradis, Claude & Strimmer 2004). Cladograms of fossil taxa are often not fully resolved, presumably owing to uncertainty in the tree topology. To deal with this, a user could individually time-scale a sample of most parsimonious trees, which are fully resolved (Bell & Braddy,inpress).Lessideally,ausercouldinputapartially unresolved consensus tree and randomly resolve the soft polytomies, generating a sample of time-scaled trees. This can be implemented within bin_timepaleophy via the argument randres and increasing ntrees. Examining multiple trees will also help us account for both the uncertainty produced by drawing taxon appearance dates from uniform distributions and the uncertainty in the phylogenetic relationships. Below is a short example of generating and plotting nine time-scaled phylogenies. timetrees <- bin_timepaleophy( cladogram,rangesdisc,add.term=t, randres=t,ntrees=9) layout(matrix(1:9,3,3)) for(i in 1:9){ plot(ladderize(timetrees[[i]]),show.tip.label=f,no.margin=t)} These time-scaled trees are very similar to each other but do differ slightly in their branch lengths and in the relationships among taxa. This variation could make a considerable difference in the results of some comparative analyses. For analyses of real data, we would want to generate samples of time-scaled trees much larger than nine to fully test our hypotheses with. The function multidiv calculates the median diversity and 95% quantile diversity across an input sample of phylogenies. Although not shown here, multidiv can also calculate diversity curves for lists, which include multiple data types, such as taxonomic range data. multidiv(timetrees) The default basic time-scaling function commonly produces zero-length branches, which are often problematic for comparative analyses (Hunt & Carrano 2010). The five time-scaling methods applicable with timepaleophy are explained in detail within the function s documentation. In addition to those methods, the paleotree library also offers the SRC time-scaling method for paleontological phylogenies (Bapst, in preparation), as utilised by the functions srctimepaleophy and bin_srctimepaleophy. To use the SRC methods, an estimate of the instantaneous sampling rate is needed. For discrete temporal data, sampling probabilities can be estimated based on the frequency distribution of taxon durations (Foote & Raup 1996). The function getsampprobdisc finds the per-interval sampling probability using models fit with maximum likelihood, using the
5 Analyses of Paleo-Trees in R 807 approach developed by Foote (1997). A similar function get- SampRateCont exists for fitting sampling models to continuous-time occurrences. getsampprobdisc also outputs an estimate of the expected completeness of the fossil record, that is, the proportion of taxa in a group that we would expect to have sampled at least once. It is very important for the likelihood optimiser of getsampprobdisc to converge, as the likelihood surface for estimating these parameters can occasionally be flat and uninformative, leading to inaccurate parameter estimates. If convergence does not occur, getsampprobdisc will print a warning message and users should refer to the paleotree documentation. #obtain sampling probability MLfit <- getsampprobdisc(rangesdisc) The sampling probability is not equivalent to the sampling rate, but can be converted using the function sprob2srate. To convert the estimate, the mean interval length must be supplied. #obtain sampling rate sprob2srate(mlfit[[2]][2],int.length=5) This estimate of the instantaneous rate can be specified as an argument in bin_srctimepaleophy. There are several important differences between the SRC algorithm and the other time-scaling methods. For example, in the SRC algorithm, polytomies in the input cladogram will always be resolved during time-scaling, terminal ranges are always added onto the phylogeny, and some taxa may be stochastically identified as ancestors (details in the paleotree documentation and Bapst in preparation). Discussion The library paleotree offers a number of functions useful for analysing or simulating phylogenetic patterns of evolution in the fossil record. A number of future improvements are planned to further extend these capabilities, such as altering getsampprobdisc to accommodate for ranges of taxa, which are still extant in the modern day. Acknowledgements I would like to particularly thank Michael Foote for comments and helping me troubleshoot several functions. I would also like to thank Matthew Pennell, Emily King, Annat Haber, Jonathan Mitchell, Graeme Lloyd and an anonymous reviewer for discussions and suggestions that have greatly improved paleotree and this manuscript. The function timepaleophy is inspired by code originally provided to me by Graeme Lloyd and Gene Hunt. References Bell, M.A. & Braddy, S.J. (in press) Cope s rule in the Ordovician trilobite family Asaphidae (order Asaphida): patterns across multiple most parsimonious trees. Historical Biology. Ezard, T.H.G. & Purvis, A. (2009) paleophylo: free software to draw paleobiological phylogenies. Paleobiology, 35, Foote, M. (1996) On the probability of ancestors in the fossil record. Paleobiology, 22, Foote, M. (1997) Estimating taxonomic durations and preservation probability. Paleobiology, 23, Foote, M. & Raup, D.M. (1996) Fossil preservation and the stratigraphic ranges of taxa. Paleobiology, 22, Fusco, G., Garland, J.T., Hunt, G. & Hughes, N.C. (2012) Developmental trait evolution in trilobites. Evolution, 66, Harmon, L.J., Weir, J.T., Brock, C.D., Glor, R.E. & Challenger, W. (2008) GEI- GER: investigating evolutionaryradiations. Bioinformatics, 24, Hopkins, M.J. (2011) How species longevity, intraspecific morphological variation, and geographic range size are related: a comparison using late cambrian trilobites. Evolution, 65, Hunt, G. & Carrano, M.T. (2010) Models and methods for analyzing phenotypic evolution in lineages and clades. Short Course on Quantitative Methods in Paleobiology (eds J. Alroy & G. Hunt), pp Paleontological Society, Denver, CO. Kendall, D.G. (1948) On the generalized "birth-and-death" process. The Annals of Mathematical Statistics, 19, Liow, L.H., Quental, T.B. & Marshall, C.R. (2010) When Can Decreasing Diversification Rates Be Detected with Molecular Phylogenies and the Fossil Record? Systematic Biology, 59, Lloyd, G.T., Wang, S.C. & Brusatte, S.L. (2012) Identifying heterogeneity in rates of morphological evolution: discrete character change in the evolution of lungfish (Sarcopterygii, Dipnoi). Evolution, 66, Nee, S. (2006) Birth-death models in macroevolution. Annual Review of Ecology, Evolution, and Systematics, 37, Paradis, E., Claude, J. & Strimmer, K. (2004) APE: analyses of phylogenetics and evolution in R language. Bioinformatics, 20, R Development Core Team (2011) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. (accessed 28 January 2012). Revell, L.J. (2012) phytools: an R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution, 3, Schliep, K.P. (2011) phangorn: phylogenetic analysis in R. Bioinformatics, 27, Smith, A.B. (1994) Systematics and the Fossil Record: Documenting Evolutionary Patterns. Blackwell Scientific, Oxford. Wagner, P.J. (1995) Stratigraphic tests of cladistic hypotheses. Paleobiology, 21, Wagner, P. & Erwin, D. (1995) Phylogenetic patterns as tests of speciation models. New Approaches to Speciation in the Fossil Record (eds D.H. Erwin & R.L. Anstey), pp Columbia University Press, New York. Received 23 February 2012; accepted 27 April 2012 Handling Editor: Robert Freckleton
Package paleotree. February 23, 2013
Package paleotree February 23, 2013 Type Package Version 1.8 Title Paleontological and Phylogenetic Analyses of Evolution Date 02-22-13 Author David Bapst Depends R (>= 2.15.0), ape Imports phangorn, geiger
More informationIntegrating Fossils into Phylogenies. Throughout the 20th century, the relationship between paleontology and evolutionary biology has been strained.
IB 200B Principals of Phylogenetic Systematics Spring 2011 Integrating Fossils into Phylogenies Throughout the 20th century, the relationship between paleontology and evolutionary biology has been strained.
More information8/23/2014. Phylogeny and the Tree of Life
Phylogeny and the Tree of Life Chapter 26 Objectives Explain the following characteristics of the Linnaean system of classification: a. binomial nomenclature b. hierarchical classification List the major
More informationIntegrative Biology 200 "PRINCIPLES OF PHYLOGENETICS" Spring 2018 University of California, Berkeley
Integrative Biology 200 "PRINCIPLES OF PHYLOGENETICS" Spring 2018 University of California, Berkeley B.D. Mishler Feb. 14, 2018. Phylogenetic trees VI: Dating in the 21st century: clocks, & calibrations;
More informationAP Biology. Cladistics
Cladistics Kingdom Summary Review slide Review slide Classification Old 5 Kingdom system Eukaryote Monera, Protists, Plants, Fungi, Animals New 3 Domain system reflects a greater understanding of evolution
More informationAmira A. AL-Hosary PhD of infectious diseases Department of Animal Medicine (Infectious Diseases) Faculty of Veterinary Medicine Assiut
Amira A. AL-Hosary PhD of infectious diseases Department of Animal Medicine (Infectious Diseases) Faculty of Veterinary Medicine Assiut University-Egypt Phylogenetic analysis Phylogenetic Basics: Biological
More informationChapter 7: Models of discrete character evolution
Chapter 7: Models of discrete character evolution pdf version R markdown to recreate analyses Biological motivation: Limblessness as a discrete trait Squamates, the clade that includes all living species
More informationC.DARWIN ( )
C.DARWIN (1809-1882) LAMARCK Each evolutionary lineage has evolved, transforming itself, from a ancestor appeared by spontaneous generation DARWIN All organisms are historically interconnected. Their relationships
More informationHow should we organize the diversity of animal life?
How should we organize the diversity of animal life? The difference between Taxonomy Linneaus, and Cladistics Darwin What are phylogenies? How do we read them? How do we estimate them? Classification (Taxonomy)
More informationDr. Amira A. AL-Hosary
Phylogenetic analysis Amira A. AL-Hosary PhD of infectious diseases Department of Animal Medicine (Infectious Diseases) Faculty of Veterinary Medicine Assiut University-Egypt Phylogenetic Basics: Biological
More informationReconstructing the history of lineages
Reconstructing the history of lineages Class outline Systematics Phylogenetic systematics Phylogenetic trees and maps Class outline Definitions Systematics Phylogenetic systematics/cladistics Systematics
More informationClassification, Phylogeny yand Evolutionary History
Classification, Phylogeny yand Evolutionary History The diversity of life is great. To communicate about it, there must be a scheme for organization. There are many species that would be difficult to organize
More informationChapter 27: Evolutionary Genetics
Chapter 27: Evolutionary Genetics Student Learning Objectives Upon completion of this chapter you should be able to: 1. Understand what the term species means to biology. 2. Recognize the various patterns
More informationHistorical Biogeography. Historical Biogeography. Systematics
Historical Biogeography I. Definitions II. Fossils: problems with fossil record why fossils are important III. Phylogeny IV. Phenetics VI. Phylogenetic Classification Disjunctions debunked: Examples VII.
More informationAnatomy of a tree. clade is group of organisms with a shared ancestor. a monophyletic group shares a single common ancestor = tapirs-rhinos-horses
Anatomy of a tree outgroup: an early branching relative of the interest groups sister taxa: taxa derived from the same recent ancestor polytomy: >2 taxa emerge from a node Anatomy of a tree clade is group
More informationMagnitude versus direction of change and the contribution of macroevolutionary trends to morphological disparity
Biological Journal of the Linnean Society, 2016, 118, 116 130. With 10 figures. Magnitude versus direction of change and the contribution of macroevolutionary trends to morphological disparity MELANIE
More informationLecture V Phylogeny and Systematics Dr. Kopeny
Delivered 1/30 and 2/1 Lecture V Phylogeny and Systematics Dr. Kopeny Lecture V How to Determine Evolutionary Relationships: Concepts in Phylogeny and Systematics Textbook Reading: pp 425-433, 435-437
More informationMacroevolution Part I: Phylogenies
Macroevolution Part I: Phylogenies Taxonomy Classification originated with Carolus Linnaeus in the 18 th century. Based on structural (outward and inward) similarities Hierarchal scheme, the largest most
More informationThe Tempo of Macroevolution: Patterns of Diversification and Extinction
The Tempo of Macroevolution: Patterns of Diversification and Extinction During the semester we have been consider various aspects parameters associated with biodiversity. Current usage stems from 1980's
More informationPhylogenies & Classifying species (AKA Cladistics & Taxonomy) What are phylogenies & cladograms? How do we read them? How do we estimate them?
Phylogenies & Classifying species (AKA Cladistics & Taxonomy) What are phylogenies & cladograms? How do we read them? How do we estimate them? Carolus Linneaus:Systema Naturae (1735) Swedish botanist &
More informationEXTINCTION RATES SHOULD NOT BE ESTIMATED FROM MOLECULAR PHYLOGENIES
ORIGINAL ARTICLE doi:10.1111/j.1558-5646.2009.00926.x EXTINCTION RATES SHOULD NOT BE ESTIMATED FROM MOLECULAR PHYLOGENIES Daniel L. Rabosky 1,2,3,4 1 Department of Ecology and Evolutionary Biology, Cornell
More informationCHAPTER 26 PHYLOGENY AND THE TREE OF LIFE Connecting Classification to Phylogeny
CHAPTER 26 PHYLOGENY AND THE TREE OF LIFE Connecting Classification to Phylogeny To trace phylogeny or the evolutionary history of life, biologists use evidence from paleontology, molecular data, comparative
More informationASYMMETRIES IN PHYLOGENETIC DIVERSIFICATION AND CHARACTER CHANGE CAN BE UNTANGLED
doi:1.1111/j.1558-5646.27.252.x ASYMMETRIES IN PHYLOGENETIC DIVERSIFICATION AND CHARACTER CHANGE CAN BE UNTANGLED Emmanuel Paradis Institut de Recherche pour le Développement, UR175 CAVIAR, GAMET BP 595,
More informationBiology 211 (2) Week 1 KEY!
Biology 211 (2) Week 1 KEY Chapter 1 KEY FIGURES: 1.2, 1.3, 1.4, 1.5, 1.6, 1.7 VOCABULARY: Adaptation: a trait that increases the fitness Cells: a developed, system bound with a thin outer layer made of
More informationChapter 19: Taxonomy, Systematics, and Phylogeny
Chapter 19: Taxonomy, Systematics, and Phylogeny AP Curriculum Alignment Chapter 19 expands on the topics of phylogenies and cladograms, which are important to Big Idea 1. In order for students to understand
More informationConcepts and Methods in Molecular Divergence Time Estimation
Concepts and Methods in Molecular Divergence Time Estimation 26 November 2012 Prashant P. Sharma American Museum of Natural History Overview 1. Why do we date trees? 2. The molecular clock 3. Local clocks
More informationIntraspecific gene genealogies: trees grafting into networks
Intraspecific gene genealogies: trees grafting into networks by David Posada & Keith A. Crandall Kessy Abarenkov Tartu, 2004 Article describes: Population genetics principles Intraspecific genetic variation
More informationClassification and Phylogeny
Classification and Phylogeny The diversity of life is great. To communicate about it, there must be a scheme for organization. There are many species that would be difficult to organize without a scheme
More informationA Rough Guide to CladeAge
A Rough Guide to CladeAge Setting fossil calibrations in BEAST 2 Michael Matschiner Remco Bouckaert michaelmatschiner@mac.com remco@cs.auckland.ac.nz January 17, 2014 Contents 1 Introduction 1 1.1 Windows...............................
More informationOrganizing Life s Diversity
17 Organizing Life s Diversity section 2 Modern Classification Classification systems have changed over time as information has increased. What You ll Learn species concepts methods to reveal phylogeny
More informationClassification and Phylogeny
Classification and Phylogeny The diversity it of life is great. To communicate about it, there must be a scheme for organization. There are many species that would be difficult to organize without a scheme
More informationAlgorithms in Bioinformatics
Algorithms in Bioinformatics Sami Khuri Department of Computer Science San José State University San José, California, USA khuri@cs.sjsu.edu www.cs.sjsu.edu/faculty/khuri Distance Methods Character Methods
More informationThe practice of naming and classifying organisms is called taxonomy.
Chapter 18 Key Idea: Biologists use taxonomic systems to organize their knowledge of organisms. These systems attempt to provide consistent ways to name and categorize organisms. The practice of naming
More informationUoN, CAS, DBSC BIOL102 lecture notes by: Dr. Mustafa A. Mansi. The Phylogenetic Systematics (Phylogeny and Systematics)
- Phylogeny? - Systematics? The Phylogenetic Systematics (Phylogeny and Systematics) - Phylogenetic systematics? Connection between phylogeny and classification. - Phylogenetic systematics informs the
More informationBAMMtools: an R package for the analysis of evolutionary dynamics on phylogenetic trees
Methods in Ecology and Evolution 2, 5, 71 77 doi: 1.1111/241-21X.12199 APPLICATION BAMMtools: an R package for the analysis of evolutionary dynamics on phylogenetic trees Daniel L. Rabosky*, Michael Grundler,
More informationPRINCIPLES OF PHYLOGENETICS: ECOLOGY AND EVOLUTION
"PRINCIPLES OF PHYLOGENETICS: ECOLOGY AND EVOLUTION" Integrative Biology 200 Spring 2018 University of California, Berkeley D. Ackerly April 2, 2018 Lineage diversification Reading: Morlon, H. (2014).
More informationThe Life System and Environmental & Evolutionary Biology II
The Life System and Environmental & Evolutionary Biology II EESC V2300y / ENVB W2002y Laboratory 1 (01/28/03) Systematics and Taxonomy 1 SYNOPSIS In this lab we will give an overview of the methodology
More informationPHYLOGENY & THE TREE OF LIFE
PHYLOGENY & THE TREE OF LIFE PREFACE In this powerpoint we learn how biologists distinguish and categorize the millions of species on earth. Early we looked at the process of evolution here we look at
More informationPhylogenetic approaches for studying diversification
Ecology Letters, (2014) doi: 10.1111/ele.12251 REVIEW AND SYNTHESIS Phylogenetic approaches for studying diversification Helene Morlon* Center for Applied Mathematics, Ecole Polytechnique, Palaiseau, Essonne,
More informationPhylogeny-Based Analyses of Evolution with a Paleo-Focus Part 2. Diversification of Lineages (Mar 02, 2012 David W. Bapst
Phylogeny-Based Analyses of Evolution with a Paleo-Focus Part 2. Diversification of Lineages (Mar 02, 2012 David W. Bapst Recommended Reading: Nee, S. 2006. Birth-Death Models in Macroevolution. Annual
More informationOutline. Classification of Living Things
Outline Classification of Living Things Chapter 20 Mader: Biology 8th Ed. Taxonomy Binomial System Species Identification Classification Categories Phylogenetic Trees Tracing Phylogeny Cladistic Systematics
More informationA Simple Method for Estimating Informative Node Age Priors for the Fossil Calibration of Molecular Divergence Time Analyses
A Simple Method for Estimating Informative Node Age Priors for the Fossil Calibration of Molecular Divergence Time Analyses Michael D. Nowak 1 *, Andrew B. Smith 2, Carl Simpson 3, Derrick J. Zwickl 4
More informationPlant of the Day Isoetes andicola
Plant of the Day Isoetes andicola Endemic to central and southern Peru Found in scattered populations above 4000 m Restricted to the edges of bogs and lakes Leaves lack stomata and so CO 2 is obtained,
More informationPhylogeny and systematics. Why are these disciplines important in evolutionary biology and how are they related to each other?
Phylogeny and systematics Why are these disciplines important in evolutionary biology and how are they related to each other? Phylogeny and systematics Phylogeny: the evolutionary history of a species
More informationPOSITIVE CORRELATION BETWEEN DIVERSIFICATION RATES AND PHENOTYPIC EVOLVABILITY CAN MIMIC PUNCTUATED EQUILIBRIUM ON MOLECULAR PHYLOGENIES
doi:10.1111/j.1558-5646.2012.01631.x POSITIVE CORRELATION BETWEEN DIVERSIFICATION RATES AND PHENOTYPIC EVOLVABILITY CAN MIMIC PUNCTUATED EQUILIBRIUM ON MOLECULAR PHYLOGENIES Daniel L. Rabosky 1,2,3 1 Department
More informationRate of Evolution Juliana Senawi
Rate of Evolution Juliana Senawi Rate of Evolution Measurement of the change in an evolutionary lineage overtime Radiometric and paleomagnetic dating provide an effective basis for determining the age
More informationIntroduction to characters and parsimony analysis
Introduction to characters and parsimony analysis Genetic Relationships Genetic relationships exist between individuals within populations These include ancestordescendent relationships and more indirect
More informationGENETICS - CLUTCH CH.22 EVOLUTIONARY GENETICS.
!! www.clutchprep.com CONCEPT: OVERVIEW OF EVOLUTION Evolution is a process through which variation in individuals makes it more likely for them to survive and reproduce There are principles to the theory
More informationC3020 Molecular Evolution. Exercises #3: Phylogenetics
C3020 Molecular Evolution Exercises #3: Phylogenetics Consider the following sequences for five taxa 1-5 and the known outgroup O, which has the ancestral states (note that sequence 3 has changed from
More informationUnit 9: Evolution Guided Reading Questions (80 pts total)
Name: AP Biology Biology, Campbell and Reece, 7th Edition Adapted from chapter reading guides originally created by Lynn Miriello Unit 9: Evolution Guided Reading Questions (80 pts total) Chapter 22 Descent
More informationChapter 22: Descent with Modification 1. BRIEFLY summarize the main points that Darwin made in The Origin of Species.
AP Biology Chapter Packet 7- Evolution Name Chapter 22: Descent with Modification 1. BRIEFLY summarize the main points that Darwin made in The Origin of Species. 2. Define the following terms: a. Natural
More informationHow to read and make phylogenetic trees Zuzana Starostová
How to read and make phylogenetic trees Zuzana Starostová How to make phylogenetic trees? Workflow: obtain DNA sequence quality check sequence alignment calculating genetic distances phylogeny estimation
More informationSTEM-hy: Species Tree Estimation using Maximum likelihood (with hybridization)
STEM-hy: Species Tree Estimation using Maximum likelihood (with hybridization) Laura Salter Kubatko Departments of Statistics and Evolution, Ecology, and Organismal Biology The Ohio State University kubatko.2@osu.edu
More informationCladistics and Bioinformatics Questions 2013
AP Biology Name Cladistics and Bioinformatics Questions 2013 1. The following table shows the percentage similarity in sequences of nucleotides from a homologous gene derived from five different species
More informationChapter 26 Phylogeny and the Tree of Life
Chapter 26 Phylogeny and the Tree of Life Biologists estimate that there are about 5 to 100 million species of organisms living on Earth today. Evidence from morphological, biochemical, and gene sequence
More informationBINF6201/8201. Molecular phylogenetic methods
BINF60/80 Molecular phylogenetic methods 0-7-06 Phylogenetics Ø According to the evolutionary theory, all life forms on this planet are related to one another by descent. Ø Traditionally, phylogenetics
More informationLecture 6 Phylogenetic Inference
Lecture 6 Phylogenetic Inference From Darwin s notebook in 1837 Charles Darwin Willi Hennig From The Origin in 1859 Cladistics Phylogenetic inference Willi Hennig, Cladistics 1. Clade, Monophyletic group,
More informationOMICS Journals are welcoming Submissions
OMICS Journals are welcoming Submissions OMICS International welcomes submissions that are original and technically so as to serve both the developing world and developed countries in the best possible
More informationELE4120 Bioinformatics Tutorial 8
ELE4120 ioinformatics Tutorial 8 ontent lassifying Organisms Systematics and Speciation Taxonomy and phylogenetics Phenetics versus cladistics Phylogenetic trees iological classification Goal: To develop
More informationUsing the Birth-Death Process to Infer Changes in the Pattern of Lineage Gain and Loss. Nathaniel Malachi Hallinan
Using the Birth-Death Process to Infer Changes in the Pattern of Lineage Gain and Loss by Nathaniel Malachi Hallinan A dissertation submitted in partial satisfaction of the requirements for the degree
More informationChapter 26: Phylogeny and the Tree of Life Phylogenies Show Evolutionary Relationships
Chapter 26: Phylogeny and the Tree of Life You Must Know The taxonomic categories and how they indicate relatedness. How systematics is used to develop phylogenetic trees. How to construct a phylogenetic
More informationPhylogeny 9/8/2014. Evolutionary Relationships. Data Supporting Phylogeny. Chapter 26
Phylogeny Chapter 26 Taxonomy Taxonomy: ordered division of organisms into categories based on a set of characteristics used to assess similarities and differences Carolus Linnaeus developed binomial nomenclature,
More informationMaximum likelihood estimation of phylogeny using stratigraphic data
Paleobiology, 23(2), 1997, pp. 174-180 Maximum likelihood estimation of phylogeny using stratigraphic data John P. Huelsenbeck and Bruce Rannala Abstract.-The stratigraphic distribution of fossil species
More informationBRIEF COMMUNICATIONS
Evolution, 59(12), 2005, pp. 2705 2710 BRIEF COMMUNICATIONS THE EFFECT OF INTRASPECIFIC SAMPLE SIZE ON TYPE I AND TYPE II ERROR RATES IN COMPARATIVE STUDIES LUKE J. HARMON 1 AND JONATHAN B. LOSOS Department
More informationvary spuriously with preservation rate, but this spurious variation is largely eliminated and true rates are reasonably well estimated.
606 MICHAEL FOOTE Figure 1 shows results of an experiment with simulated data. Rates of origination, extinction, and preservation were varied in a pulsed pattern, in which a background level was punctuated
More informationBiology 559R: Introduction to Phylogenetic Comparative Methods Topics for this week:
Biology 559R: Introduction to Phylogenetic Comparative Methods Topics for this week: Course general information About the course Course objectives Comparative methods: An overview R as language: uses and
More informationUsing phylogenetics to estimate species divergence times... Basics and basic issues for Bayesian inference of divergence times (plus some digression)
Using phylogenetics to estimate species divergence times... More accurately... Basics and basic issues for Bayesian inference of divergence times (plus some digression) "A comparison of the structures
More informationNon-independence in Statistical Tests for Discrete Cross-species Data
J. theor. Biol. (1997) 188, 507514 Non-independence in Statistical Tests for Discrete Cross-species Data ALAN GRAFEN* AND MARK RIDLEY * St. John s College, Oxford OX1 3JP, and the Department of Zoology,
More information1/27/2010. Systematics and Phylogenetics of the. An Introduction. Taxonomy and Systematics
Systematics and Phylogenetics of the Amphibia: An Introduction Taxonomy and Systematics Taxonomy, the science of describing biodiversity, mainly naming unnamed species, and arranging the diversity into
More informationPackage WGDgc. June 3, 2014
Package WGDgc June 3, 2014 Type Package Title Whole genome duplication detection using gene counts Version 1.1 Date 2014-06-03 Author Tram Ta, Charles-Elie Rabier, Cecile Ane Maintainer Tram Ta
More informationPackage WGDgc. October 23, 2015
Package WGDgc October 23, 2015 Version 1.2 Date 2015-10-22 Title Whole genome duplication detection using gene counts Depends R (>= 3.0.1), phylobase, phyext, ape Detection of whole genome duplications
More information"PRINCIPLES OF PHYLOGENETICS: ECOLOGY AND EVOLUTION" Integrative Biology 200B Spring 2011 University of California, Berkeley
"PRINCIPLES OF PHYLOGENETICS: ECOLOGY AND EVOLUTION" Integrative Biology 200B Spring 2011 University of California, Berkeley B.D. Mishler March 31, 2011. Reticulation,"Phylogeography," and Population Biology:
More informationA (short) introduction to phylogenetics
A (short) introduction to phylogenetics Thibaut Jombart, Marie-Pauline Beugin MRC Centre for Outbreak Analysis and Modelling Imperial College London Genetic data analysis with PR Statistics, Millport Field
More informationBiology 2. Lecture Material. For. Macroevolution. Systematics
Biology 2 Macroevolution & Systematics 1 Biology 2 Lecture Material For Macroevolution & Systematics Biology 2 Macroevolution & Systematics 2 Microevolution: Biological Species: Two Patterns of Evolutionary
More informationTowards a general model of superorganism life history
Figure S1 Results from the scaling of phylogenetically independent contrasts (PIC). For each analysis, we provide the tree used to calculate contrasts as well as the OLS regression used to calculate scaling
More informationBiology. Slide 1 of 24. End Show. Copyright Pearson Prentice Hall
Biology 1 of 24 18-2 Modern Evolutionary Classification 2 of 24 18-2 Modern Evolutionary Classification Evolutionary Classification Evolutionary Classification Phylogeny is the study of evolutionary relationships
More informationEstimating Age-Dependent Extinction: Contrasting Evidence from Fossils and Phylogenies
Syst. Biol. 0(0):1 17, 2017 The Author(s) 2017. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. This is an Open Access article distributed under the terms of the
More informationRapid evolution of the cerebellum in humans and other great apes
Rapid evolution of the cerebellum in humans and other great apes Article Accepted Version Barton, R. A. and Venditti, C. (2014) Rapid evolution of the cerebellum in humans and other great apes. Current
More informationESS 345 Ichthyology. Systematic Ichthyology Part II Not in Book
ESS 345 Ichthyology Systematic Ichthyology Part II Not in Book Thought for today: Now, here, you see, it takes all the running you can do, to keep in the same place. If you want to get somewhere else,
More informationAppendix from L. J. Revell, On the Analysis of Evolutionary Change along Single Branches in a Phylogeny
008 by The University of Chicago. All rights reserved.doi: 10.1086/588078 Appendix from L. J. Revell, On the Analysis of Evolutionary Change along Single Branches in a Phylogeny (Am. Nat., vol. 17, no.
More informationBiol 206/306 Advanced Biostatistics Lab 11 Models of Trait Evolution Fall 2016
Biol 206/306 Advanced Biostatistics Lab 11 Models of Trait Evolution Fall 2016 By Philip J. Bergmann 0. Laboratory Objectives 1. Explore how evolutionary trait modeling can reveal different information
More informationInferring the Dynamics of Diversification: A Coalescent Approach
Inferring the Dynamics of Diversification: A Coalescent Approach Hélène Morlon 1 *, Matthew D. Potts 2, Joshua B. Plotkin 1 * 1 Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania,
More informationPyRate: a new program to estimate speciation and extinction rates from incomplete fossil data
Methods in Ecology and Evolution 2014, 5, 1126 1131 doi: 10.1111/2041-210X.12263 APPLICATION PyRate: a new program to estimate speciation and extinction rates from incomplete fossil data Daniele Silvestro
More informationAn introduction to the picante package
An introduction to the picante package Steven Kembel (steve.kembel@gmail.com) April 2010 Contents 1 Installing picante 1 2 Data formats in picante 2 2.1 Phylogenies................................ 2 2.2
More informationAPPENDIX S1: DESCRIPTION OF THE ESTIMATION OF THE VARIANCES OF OUR MAXIMUM LIKELIHOOD ESTIMATORS
APPENDIX S1: DESCRIPTION OF THE ESTIMATION OF THE VARIANCES OF OUR MAXIMUM LIKELIHOOD ESTIMATORS For large n, the negative inverse of the second derivative of the likelihood function at the optimum can
More informationThe terrestrial rock record
The terrestrial rock record Stratigraphy, vertebrate biostratigraphy and phylogenetics The Cretaceous-Paleogene boundary at Hell Creek, Montana. Hell Creek Fm. lower, Tullock Fm. upper. (P. David Polly,
More informationPhylogeny: building the tree of life
Phylogeny: building the tree of life Dr. Fayyaz ul Amir Afsar Minhas Department of Computer and Information Sciences Pakistan Institute of Engineering & Applied Sciences PO Nilore, Islamabad, Pakistan
More informationDoes convergent evolution always result from different lineages experiencing similar
Different evolutionary dynamics led to the convergence of clinging performance in lizard toepads Sarin Tiatragula 1 *, Gopal Murali 2 *, James T. Stroud 3 1 Department of Biological Sciences, Auburn University,
More informationPackage FossilSim. R topics documented: November 16, Type Package Title Simulation of Fossil and Taxonomy Data Version 2.1.0
Type Package Title Simulation of Fossil and Taxonomy Data Version 2.1.0 Package FossilSim November 16, 2018 Simulating taxonomy and fossil data on phylogenetic s under mechanistic models of speciation,
More informationIntegrative Biology 200A "PRINCIPLES OF PHYLOGENETICS" Spring 2012 University of California, Berkeley
Integrative Biology 200A "PRINCIPLES OF PHYLOGENETICS" Spring 2012 University of California, Berkeley B.D. Mishler April 12, 2012. Phylogenetic trees IX: Below the "species level;" phylogeography; dealing
More informationMarkov chain Monte-Carlo to estimate speciation and extinction rates: making use of the forest hidden behind the (phylogenetic) tree
Markov chain Monte-Carlo to estimate speciation and extinction rates: making use of the forest hidden behind the (phylogenetic) tree Nicolas Salamin Department of Ecology and Evolution University of Lausanne
More informationProcesses of Evolution
15 Processes of Evolution Forces of Evolution Concept 15.4 Selection Can Be Stabilizing, Directional, or Disruptive Natural selection can act on quantitative traits in three ways: Stabilizing selection
More informationEvolutionary trends. Horse size increased steadily. Phylogeny and the fossil record
Phylogeny and the fossil record The fossil record: trends and rates Chapter 4 Strong correspondence between phylogenetic branching order and order of appearance in the fossil record Evolutionary trends
More informationIntegrative Biology 200A "PRINCIPLES OF PHYLOGENETICS" Spring 2012 University of California, Berkeley
Integrative Biology 200A "PRINCIPLES OF PHYLOGENETICS" Spring 2012 University of California, Berkeley B.D. Mishler Feb. 7, 2012. Morphological data IV -- ontogeny & structure of plants The last frontier
More informationHominid Evolution What derived characteristics differentiate members of the Family Hominidae and how are they related?
Hominid Evolution What derived characteristics differentiate members of the Family Hominidae and how are they related? Introduction. The central idea of biological evolution is that all life on Earth shares
More informationChapter 16: Reconstructing and Using Phylogenies
Chapter Review 1. Use the phylogenetic tree shown at the right to complete the following. a. Explain how many clades are indicated: Three: (1) chimpanzee/human, (2) chimpanzee/ human/gorilla, and (3)chimpanzee/human/
More informationEstimating a Binary Character s Effect on Speciation and Extinction
Syst. Biol. 56(5):701 710, 2007 Copyright c Society of Systematic Biologists ISSN: 1063-5157 print / 1076-836X online DOI: 10.1080/10635150701607033 Estimating a Binary Character s Effect on Speciation
More informationThree Monte Carlo Models. of Faunal Evolution PUBLISHED BY NATURAL HISTORY THE AMERICAN MUSEUM SYDNEY ANDERSON AND CHARLES S.
AMERICAN MUSEUM Notltates PUBLISHED BY THE AMERICAN MUSEUM NATURAL HISTORY OF CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2563 JANUARY 29, 1975 SYDNEY ANDERSON AND CHARLES S. ANDERSON
More informationBiology 1B Evolution Lecture 2 (February 26, 2010) Natural Selection, Phylogenies
1 Natural Selection (Darwin-Wallace): There are three conditions for natural selection: 1. Variation: Individuals within a population have different characteristics/traits (or phenotypes). 2. Inheritance:
More informationBioinformatics tools for phylogeny and visualization. Yanbin Yin
Bioinformatics tools for phylogeny and visualization Yanbin Yin 1 Homework assignment 5 1. Take the MAFFT alignment http://cys.bios.niu.edu/yyin/teach/pbb/purdue.cellwall.list.lignin.f a.aln as input and
More information