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Donoghue Evolution of disparity - University of Bristol · range of designs (morphological...

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Patterns, processes, rates, and constraints, in the evolution of morphological disparity Supervisors: Philip Donoghue (Bristol), Matthew Wills (Bath), Davide Pisani (Bristol), Mario dos Reis (Queen Mary, London), Thomas Guillerme (Imperial) From Aristotle onwards, it has been apparent that there are discontinuities in organismal form, such that species are clumped within morphological ‘design space’ [1]. These clumps at once help us in attempts to classify but often frustrate our efforts to resolve phylogeny. There is vigorous debate as to why organisms are irregularly distributed in this way. Do clusters reflect peaks in a rugged fitness landscape? Are the intervening regions of design space impossible to colonise – either because of physical or developmental constraints – or are they unoccupied because insufficient time has elapsed? Were they once occupied and subsequently vacated in the wake of past extinctions? Is the range of designs (morphological disparity) realised progressively through time, or in an early burst of innovation followed by stasis? The answers to these questions have important implications, particularly whether the same processes shape macroevolution throughout deep time (uniformitarianism) or whether these forces change [2]. To date, analyses have been based largely on empirical case studies of extinct organisms. Little attempt has been made to establish a theoretical or empirical foundation for interpreting empirical disparity analyses. This is unsurprising, since methods for simulating the evolution of morphological data have only recently become available [3]. This project will augment these tools in order to simulate phenotypic evolution under the diversity of evolutionary models that are inferred to have shaped the morphological disparity of real clades. These include intrinsic factors (such as changes in rates of character evolution and the scale of innovation, phylogenetic burden, and lineage duration) as well as extrinsic factors (such as Figure: Competing models for the evolution of morphological disparity - from [4]
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Page 1: Donoghue Evolution of disparity - University of Bristol · range of designs (morphological disparity) realised progressively through time, or in an early burst of innovation followed

Patterns,processes,rates,andconstraints,intheevolutionofmorphologicaldisparitySupervisors:PhilipDonoghue(Bristol),MatthewWills(Bath),DavidePisani(Bristol),MariodosReis(QueenMary,London),ThomasGuillerme(Imperial)FromAristotleonwards,ithasbeenapparentthattherearediscontinuitiesinorganismalform,suchthatspeciesareclumpedwithinmorphological‘designspace’[1].Theseclumpsatoncehelpusinattemptstoclassifybutoftenfrustrateoureffortstoresolvephylogeny.Thereisvigorousdebateastowhyorganismsareirregularlydistributedinthisway.Doclustersreflectpeaksinaruggedfitnesslandscape?Aretheinterveningregionsofdesignspaceimpossibletocolonise–eitherbecauseofphysicalordevelopmentalconstraints–oraretheyunoccupiedbecauseinsufficienttimehaselapsed?Weretheyonceoccupiedandsubsequentlyvacatedinthewakeofpastextinctions?Istherangeofdesigns(morphologicaldisparity)realisedprogressivelythroughtime,orinanearlyburstofinnovationfollowedbystasis?Theanswerstothesequestionshaveimportantimplications,particularlywhetherthesameprocessesshapemacroevolutionthroughoutdeeptime(uniformitarianism)orwhethertheseforceschange[2].

Todate,analyseshavebeenbasedlargelyonempiricalcasestudiesofextinctorganisms.Littleattempthasbeenmadetoestablishatheoreticalorempiricalfoundationforinterpretingempiricaldisparityanalyses.Thisisunsurprising,sincemethodsforsimulatingtheevolutionofmorphologicaldatahaveonlyrecentlybecomeavailable[3].Thisprojectwillaugmentthesetoolsinordertosimulatephenotypicevolutionunderthediversityofevolutionarymodelsthatareinferredtohaveshapedthemorphologicaldisparityofrealclades.Theseincludeintrinsicfactors(suchaschangesinratesofcharacterevolutionandthescaleofinnovation,phylogeneticburden,andlineageduration)aswellasextrinsicfactors(suchas

Figure:Competingmodelsfortheevolutionofmorphologicaldisparity-from[4]

Page 2: Donoghue Evolution of disparity - University of Bristol · range of designs (morphological disparity) realised progressively through time, or in an early burst of innovation followed

competitionandmassextinctionevents).Theseputativedriversandlimitswillbeexaminedinturnandincombinationtodeterminewhethertheyyieldthepatternsofdesignspaceoccupationthatareobservedinrealclades.Thecandidatewillalsoexplorehowthestructuralandphylogeneticrelationshipsamongcharacterscontributestomorphospaceoccupation.WhiletheintentionistosamplebroadlyacrossorganismaldiversityandthroughoutthePhanerozoic.thereisalsoscopetofocusonparticulargroups,dependinguponthesuccessfulcandidate’sinterest.Ultimately,theresultsofthesimulationstudieswillidentifytherelativerolesofdifferentevolutionaryprocessesinshapingtheoccupationofdesignspace,andprovideanewtheoreticalframeworkwithinwhichtoevaluatetheresultsofempiricaldisparityanalyses.Theywillalsospeaktoaseminalcontroversyinmacroevolution,namelytheuniformitarian/non-uniformitariandebate.References1. D.H.Erwin,Disparity:morphologicpatternanddevelopmentalcontext.Palaeontology50,57-73(2007).2. D.H.Erwin,Evolutionaryuniformitarianism.Developmentalbiology357,27-34(2011).3. J.E.O'Reillyetal.,Bayesianmethodsoutperformparsimonybutattheexpenseofprecisionintheestimationofphylogenyfromdiscretemorphologicaldata.BiologyLetters12,20160081(2016).4. J.W.Oyston,M.Hughes,S.Gerber,M.A.Wills,Whyshouldweinvestigatethemorphologicaldisparityofplantclades?Annalsofbotany,(2015).


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