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    rac ng pec es n pacerac ng pec es n pace

    Assessing the relationshipsAssessing the relationships

    ,

    paleoecology, and macroevolution

    ,

    paleoecology, and macroevolution

    Al cia L. Sti allAl cia L. Sti all

    Department of Geological Sciences andDepartment of Geological Sciences and

    OHIO Center for Ecology and Evolutionary StudiesOHIO Center for Ecology and Evolutionary Studies

    Department of Geological Sciences andDepartment of Geological Sciences and

    OHIO Center for Ecology and Evolutionary StudiesOHIO Center for Ecology and Evolutionary Studies

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    OutlineOutline

    New methods and emerging opportunities inNew methods and emerging opportunities in

    Biogeographic controls on macroevolutionBiogeographic controls on macroevolution

    Biogeographic range and paleoecology/extinctionBiogeographic range and paleoecology/extinction

    Potential for synthesisPotential for synthesis

    Miocene Radiation ofMiocene Radiation of EquinaeEquinae

    Late Ordovician Richmondian InvasionLate Ordovician Richmondian Invasion

    Future research directionsFuture research directions

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    Controls on species rangeControls on species range

    Historical FactorsHistorical Factors

    Primar durin s eciationPrimar durin s eciation

    1 4 1 ,3 1,4,5 1 1 1 1 1 1 1 2 6 3 2

    1 1 1 21 1 2

    11

    Ecological FactorsEcological Factors

    1

    Primary during speciesPrimary during species

    duration & extinctionduration & extinction

    1

    1,4

    Traditionally separateTraditionally separate

    subdiciplinessubdiciplines

    Historical BiogeographyHistorical Biogeography

    Ecological BiogeographyEcological Biogeography

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    Integrated biogeographic approach, part IIntegrated biogeographic approach, part I

    Relationship between biogeography & macroevolutionRelationship between biogeography & macroevolution

    Research uestions:Research uestions:

    Speciation modeSpeciation mode CladeClade historyhistory

    Adaptive radiationsAdaptive radiations

    TectonicTectonic vsvs climatic influencesclimatic influences

    ,,

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    Phylogenetic biogeographyPhylogenetic biogeography

    Evolutionary relationshipsEvolutionary relationships Evolution of biogeographic areasEvolution of biogeographic areas

    1,2 1 6 4 1 1 1,3 1,6 3 1 3 3 4 4,6

    1,4,6 1

    1 31,3

    3

    33,4

    Areas of Endemism:Areas of Endemism:Areas of Endemism:Areas of Endemism:

    1

    1

    1

    1 1 Northern Appalachian basin1 Northern Appalachian basin

    2 Southern Appalachian basin2 Southern Appalachian basin

    3 Michigan Basin3 Michigan Basin

    4 Iowa Basin4 Iowa Basin

    5 Missouri Basin5 Missouri Basin

    6 Western United States6 Western United States

    1 Northern Appalachian basin1 Northern Appalachian basin

    2 Southern Appalachian basin2 Southern Appalachian basin

    3 Michigan Basin3 Michigan Basin

    4 Iowa Basin4 Iowa Basin

    5 Missouri Basin5 Missouri Basin

    6 Western United States6 Western United States

    1

    S eciation modeS eciation mode Relationshi of bio eo ra hic areasRelationshi of bio eo ra hic areas1,2 1 6 4 1 1 1,3 1,6 3 1 3 3 4 4,6

    1,4,6 1

    1

    1,31,3

    3

    33,4

    Ancestor

    Bolivia

    Europe

    W. Canada

    Ancestor

    Bolivia

    Europe

    W. Canada

    1

    1

    1

    1

    1 Michigan B.

    Appalachian B.

    Western US

    Central US

    Michigan B.

    Western US

    Appalachian B.

    Central US

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    Example 1: MioceneExample 1: Miocene EquinaeEquinae

    Maguire & Stigall (2008), Palaeo3

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    EquidEquid PhylogenticPhylogentic BiogeographyBiogeography

    Speciation modeSpeciation mode

    mode of speciationmode of speciation

    Dispersal

    yc ca processesyc ca processesdrive evolution ofdrive evolution of

    area relationshipsarea relationships

    Vicariance vs. GeodispersalVicariance vs. GeodispersalMaguire & Stigall (2008), Palaeo3

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    Example 2: Late Devonian Biodiversity CrisisExample 2: Late Devonian Biodiversity Crisis

    Species invasions due toSpecies invasions due to

    continental assembly andcontinental assembly and

    trangress onstrangress ons resu te nresu te n

    greatly reduced faunalgreatly reduced faunal

    endemismendemism

    Extinction rates elevatedExtinction rates elevated

    Speciation ratesSpeciation rates

    North AmericaNorth America Europe N. AfricaEurope N. Africa NorthernNorthern

    depresseddepressed

    Cosmopolitan FaunaCosmopolitan FaunaLateLate

    DevonianDevonian

    Basin/OldBasin/Old

    WorldWorld

    MichiganMichigan ENAENA Old WorldOld World

    VenezuelanVenezuelan--

    ColumbianColumbian

    MiddleMiddle

    DevonianDevonian

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    Late Devonian Speciation ModeLate Devonian Speciation Mode1,2 1 6 4 1 1 1,3 1,6 3 1 3 3 4 4,6

    1,4,6 1

    1,31,3

    3

    33,4

    0 1 4 4 3 3,4 2 3,4 2 1 3 4 1 1,3,4 3 0 1,2,4 1,3,4 5 2 1,3,4,5

    44

    4

    4

    4 44

    4

    44

    3,4

    1

    1

    1

    1

    1

    episode of vicarianceepisode of dispersal (invasion)

    episode of vicarianceepisode of dispersal (invasion)

    4

    44

    4

    4 4

    44

    AreaArea cladogramcladogram fromfrom

    Sti all Rode 2005Sti all Rode 2005AreaArea cladogramcladogram fromfrom

    1

    5200 0 000000

    0

    0

    0

    0 00 01 1 13 544 2 7

    00

    0 0

    0

    000

    2,8

    0,1,47,9 0 00 0 0 00 1 1 13 556

    00

    00

    0

    00

    0

    00

    00

    0

    11

    0,5 0,1

    1 4 1 ,3 1,4,5 1 1 1 1 1 1 1 2 6 3 2

    1 1 1 21 1 2

    10

    0

    0

    0

    0 1

    1

    00,3AreaArea cladogramcladogram fromfrom

    Rode & Lieberman (2002)Rode & Lieberman (2002)

    Overall,LOW rates of vicariance,

    but rampant dispersal

    1

    1,4

    AreaArea cladogramcladogram

    from Rode (2004)from Rode (2004)

    Low vicariance due to lack of opportunities for isolation duringLow vicariance due to lack of opportunities for isolation duringinvasive regime & contributed to speciation rate declineinvasive regime & contributed to speciation rate decline

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    Late Devonian Speciation ModeLate Devonian Speciation Mode

    Speciation by vicariance virtually nonSpeciation by vicariance virtually non--existentexistent

    spec a on v a spersa .e. spec es nvas onsspec a on v a spersa .e. spec es nvas ons

    Number of Number of Percent Percent

    events

    events

    vicariance

    by dispersal

    Schizophoria (Schizophoria) 2 17 11% 89%

    ower a

    Leptodesma (Leiopteria) 2 6 25% 75%

    Archaeostraca 6 13 32% 68%

    Overall 17 43 28% 72%

    Modern Fauna 70% 30%

    , ., .

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    Late Devonian Speciation RateLate Devonian Speciation Rate

    Extinction not elevated above background during Late DevonianExtinction not elevated above background during Late Devonian

    Speciation rate declines to near zeroSpeciation rate declines to near zero Stigall (2008)Stigall (2008)

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    Late Devonian Speciation SummaryLate Devonian Speciation Summary

    During the Late Devonian crisis interval:During the Late Devonian crisis interval:

    S eciation rates a roach zeroS eciation rates a roach zero

    Extinction elevated, but not in excess of background ratesExtinction elevated, but not in excess of background rates

    --

    SpeciationSpeciation byby vicariancevicariance virtually nonvirtually non--existentexistent

    All speciation via dispersal (i.e. species invasionsAll speciation via dispersal (i.e. species invasions))

    Breakdown ofBreakdown of biogeographicbiogeographic andandpaleoecologicpaleoecologic barriers resulted inbarriers resulted in

    s u own os u own o a opa r ca opa r c spec a onspec a on

    Rode & Lieberman (2004), PalaeoRode & Lieberman (2004), Palaeo33

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    Integrated biogeographic approach, part IIIntegrated biogeographic approach, part II

    Relationship between biogeography &Relationship between biogeography &

    paleoecologypaleoecology

    Research questionsResearch questions Range contractions andRange contractions and

    Niche evolution vs. nicheNiche evolution vs. niche

    constancyconstancy

    22

    Habitat tracking of communities vs.Habitat tracking of communities vs.

    individualistic species responseindividualistic species response

    --

    incorporating environmental variablesincorporating environmental variables

    Provides data amenable toProvides data amenable to

    testingtesting

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    Methods of quantitative range modelingMethods of quantitative range modeling

    GIS based approachesGIS based approachesPraewaagenoconchaPraewaagenoconcha speciosaspeciosa

    Polygon enclosure modelsPolygon enclosure models

    Requires robust set ofRequires robust set of

    Create minimum convexCreate minimum convex

    hullshulls 17,000 km17,000 km22

    Ecological niche modelsEcological niche models

    Requires (1) robust set ofRequires (1) robust set of

    spec es occurrence a aspec es occurrence a aAND (2) robust set ofAND (2) robust set of

    environmental parametersenvironmental parameters

    determined fromdetermined from

    sedimentologicalsedimentological proxiesproxies53,500 km53,500 km22

    Stigall Rode & Lieberman (2005)

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    Environmental data and interpolationEnvironmental data and interpolation

    Ecological Niche ModelsEcological Niche Models

    Predict species ranges to occupy the geographicPredict species ranges to occupy the geographic

    extent of the fundamental nicheextent of the fundamental niche

    Utilize computer learning based (genetic) algorithm toUtilize computer learning based (genetic) algorithm to

    known occurrence sites and environmental dataknown occurrence sites and environmental data

    Water de thWater de th Le todesma s ineri umLe todesma s ineri um

    100 0 200100 0 200

    Data collectionData collectionEnvironmental interpolationEnvironmental interpolation

    Range predict ionRange predict ion

    Stigall Rode & Lieberman (2005)

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    Example 2: Late Devonian Biodiversity CrisisExample 2: Late Devonian Biodiversity Crisis

    GISGIS--based geographicbased geographic

    range reconstructionrange reconstruction,,

    2000; detailed in Stigall, 2006)2000; detailed in Stigall, 2006)

    Over 5000 data points usedOver 5000 data points used

    to reconstruct ranges of 341to reconstruct ranges of 341

    species in 19 temporal binsspecies in 19 temporal bins(Rode & Lieberman, 2004)(Rode & Lieberman, 2004)

    ######

    #

    #

    #

    ######

    ###

    ##

    #

    # #

    ###

    ### ##### Range ofRange of TylothyrisTylothyris mesacostalismesacostalis

    FrasnianFrasnian): 10,309 km): 10,309 km22

    Rode & Lieberman (2004), Palaeo3

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    Example 2: Late Devonian Biodiversity CrisisExample 2: Late Devonian Biodiversity Crisis

    ENM analysisENM analysis

    factors:factors:Percent mud, silt, sandPercent mud, silt, sand Ox enationOx enation Substrate t eSubstrate t e

    Percent limestonePercent limestone IchnofaciesIchnofacies BiofaciesBiofacies

    Depositional environmentDepositional environment Bedding styleBedding style Water depthWater depth

    Inferred water depthInferred water depth

    Inferred water depthInferred water depthPercent mudPercent mudPercent siltPercent silt

    Stigall Rode & Lieberman (2005)

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    Temporal range changeTemporal range changePraewaagenoconchaPraewaagenoconcha speciosaspeciosa CariniferellaCariniferella carinatacarinata

    (Late varcus zone)(Late varcus zone)punctatapunctata ZoneZone LateLate varcusvarcus ZoneZone

    linguiformislinguiformis ZoneZone linguiformislinguiformis ZoneZone

    Stigall Rode & Lieberman (2005)

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    IdentifyIdentify interbasinalinterbasinal species invasionsspecies invasions

    Example:Example: PseudatrypaPseudatrypa devonianadevoniana

    Two invasion events:Two invasion events:

    (mid(mid FrasnianFrasnian, onset of TR cycle, onset of TR cycle IIcIIc))

    2)2) Iowa to New Mexico basins inIowa to New Mexico basins in rhenanarhenana zonezone(Late(Late FrasnianFrasnian, onset of TR cycle, onset of TR cycle IIdIId))

    TotalTotal interbasinalinterbasinal invasionsinvasions

    11

    Rode & Lieberman (2004), Palaeo3Stigall & Lieberman (2006), J Biogeo

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    Examine survival vs. geographic rangeExamine survival vs. geographic range

    1. Comparison of geographic range size vs. survival1. Comparison of geographic range size vs. survival

    VictimsVictims SurvivorsSurvivors

    N=30N=30

    Mean range: 6212Mean range: 6212

    SE mean: 895SE mean: 895

    N=127N=127

    Mean range: 15446Mean range: 15446

    SE mean: 3592SE mean: 3592

    2. Com arison of survival status vs. invasive histor2. Com arison of survival status vs. invasive histor

    TT--test: Htest: Hoo:: ss >> vv N=157N=157 p= 0.009p= 0.009

    X2 table

    Invasive speciesInvasive species NonNon--invasive speciesinvasive species

    ExtinctExtinct 181827.8627.86

    10910999.1499.14

    SurviveSurvive1616 1212

    N= 155N= 155 pEE

    .. ..

    Rode & Lieberman (2004), Palaeo3

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    Integrated biogeographic approach IIIIntegrated biogeographic approach III

    Complex feedback loops between biogeography,Complex feedback loops between biogeography,

    paleoecology, and macroevolutionpaleoecology, and macroevolution

    Research questions:Research questions: Biodiversity crisesBiodiversity crises

    n er as nan er as na nvas on o c n erc ange even snvas on o c n erc ange even s

    Tools: integrated analyses of phylogenetic andTools: integrated analyses of phylogenetic and

    environmental patternsenvironmental patterns

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    Case Study 3:Case Study 3: RichmondianRichmondian InvasionInvasion

    Late Ordovician invasion ofLate Ordovician invasion of taxataxa fromfrom

    W. North America into ENAW. North America into ENA

    Related to oceanographic changesRelated to oceanographic changes(Holland &(Holland & PatzkowskyPatzkowsky, 1997), 1997)

    (Holland &(Holland & PatzkowskyPatzkowsky, 2007), 2007)

    Stratigraphic frameworkStratigraphic framework(after Holland &(after Holland & PatzkowskyPatzkowsky, 1996), 1996)

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    GISGIS--based geographic range reconstructionbased geographic range reconstruction

    Geographic range ofGeographic range ofEntire speciesEntire species

    occurrence data setoccurrence data sete erte ae erte a occ enta socc enta s

    C4C4 C5C5 C6C6

    C1C1 C2C2 C3C3

    Stigall & Smith, in reviewStigall & Smith, in review

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    StratigraphicStratigraphic distribution of speciesdistribution of species

    Native species:Native species:

    Species C1 C2 C3 C4 C5 C6

    Dalmanella multisecta

    Leptaenagibbosa

    Platystrophia auburnensis

    Platystrophia corryvillensis

    Platystrophiacrassa

    Platystrophia hopensis

    Platystrophia morrowensis

    RestrictedRestricted toto

    MaysvillianMaysvillian

    Platystrophia sublaticosta

    Plectorthis aequivalis

    Plectorthis fissicosta

    Plectorthis neglecta

    Plectorthis plicatella

    Sowerbyellarugosa

    Strophomenamaysvillensis

    Strophomenaplanoconvexa

    Native species:Native species:

    Carryover toCarryover to

    RichmondianRichmondian

    ygosp rac nc nna ens s

    Dalmanellameeki

    Hebertellaoccidentalis

    Hebertellasubjugata

    Platystrophiacypha

    Platystrophialaticosta

    Platystrophiaponderosa

    Rafinesquina

    alternata

    Zygospira modesta

    DescendantsDescendants ofof

    native species:native species:

    SpeciateSpeciate inin

    Hebertella alveata

    Platystrophia acutilirata

    Platystrophiaannieana

    Platystrophiaclarksvillensis

    Platystrophiacummingsi

    Platystrophiaelkhornensis

    Platystrophiaforestei

    Platystrophiamoritura

    RichmondianRichmondian Strop omenaconcor ens s

    Strophomena nutans

    Strophomena planumbona

    Strophomena sulcata

    Strophomena vetusta

    Austinella scovellei

    Catazyga schuchertana

    Eochonetes clarksvillensis

    Gl torthis inscul ta

    RichmondianRichmondian

    extrabasinalextrabasinal

    invadersinvaders

    Hiscobeccuscapax

    Holtedahlinasulcata

    Lepidocyclusperlamellosum

    Leptaenarichmondensis

    Plaesiomys

    subquadrata

    Retrorsirostra carleyi

    Rhynchotrema denatum

    Tetraphalerella neglecta

    Stigall & Smith, inStigall & Smith, inreviewreview

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    Native species vs.Native species vs. extrabasinalextrabasinal invadersinvaders

    All Native SpeciesAll Native Species

    1. Comparison of geographic range of native vs. invasive species1. Comparison of geographic range of native vs. invasive species

    TT--test: N = 59test: N = 59

    12001200 2400240018001800 3000300095% CI for mean geographic range (km95% CI for mean geographic range (km22))

    . .

    MaysvillianMaysvillian onlyonly

    CarryoverCarryover TaxaTaxa

    TT--test: N = 39test: N = 39

    p < 0.0005p < 0.0005

    . ompar son o surv va s a us o na ve spec es vs. geograp c range. ompar son o surv va s a us o na ve spec es vs. geograp c range

    00 2400240012001200 36003600

    95% CI for mean geographic range (km95% CI for mean geographic range (km22))

    3. Comparison of species groups vs. geographic range3. Comparison of species groups vs. geographic range

    ANOVA: N = 59ANOVA: N = 59

    p = 0.025p = 0.025

    MaysvillianMaysvillian RestrictedRestricted

    ExtrabasinalExtrabasinal InvadersInvaders

    CarryoverCarryover TaxaTaxa

    00 2400240012001200 3600360095% CI for mean geographic range (km95% CI for mean geographic range (km22))

    RichmondianRichmondian SpeciationSpeciation

    Stigall, 2008Stigall, 2008

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    HebertellaHebertella phylogeny & biogeographyphylogeny & biogeography

    Geographic

    Area

    MidcontinentSpeciation

    style

    Nashville

    Dome

    Northern

    Canada

    Dispersal

    (Invasion

    Events)

    Vicariance

    KlingensmithKlingensmith (2007)(2007)

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    Integrating historical and ecologicalIntegrating historical and ecological

    pa eo ogeograp ypa eo ogeograp y

    feedbacks often masked in single approach analysesfeedbacks often masked in single approach analyses

    In the Late Devonian and Late Ordovician caseIn the Late Devonian and Late Ordovician case

    studies:studies:

    ranges (=ecological generalists)ranges (=ecological generalists)

    Preferential survival of invader taxa (typicallyPreferential survival of invader taxa (typically

    ecological generalists)ecological generalists) Reduced opportunities for vicarianceReduced opportunities for vicariance

    Increased invasions result in decreased speciationIncreased invasions result in decreased speciation

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    ConclusionsConclusions

    Quantitative methods provide new analytical rigor toQuantitative methods provide new analytical rigor topaleobiogeographypaleobiogeography

    Potential to analyze complexPotential to analyze complex paleobiologicalpaleobiological patternspatterns

    Capacity for hypothesis testing and generationCapacity for hypothesis testing and generation

    Emerging research questionsEmerging research questions

    Relationship between species ranges and speciationRelationship between species ranges and speciation Range expansion and contraction under shiftingRange expansion and contraction under shifting

    paleoecologicalpaleoecological regimesregimes

    Impact of invasive species on community structure andImpact of invasive species on community structure and

    macroevolutionarymacroevolutionary dynamicsdynamics Mechanics of transitions between endemic andMechanics of transitions between endemic and

    cosmopolitan faunascosmopolitan faunas

    How ecology and geographic range impact extinctionHow ecology and geographic range impact extinction

    during both background and crisis intervalsduring both background and crisis intervals

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    AcknowledgementsAcknowledgements

    Collaborators:Collaborators: Bruce LiebermanBruce Lieberman

    Students:Students: B.B. KlingensmithKlingensmith, K. Maguire, J. Smith, K. Maguire, J. Smith

    Funding:Funding: Ohio University, ACSs PetroleumOhio University, ACSs Petroleum

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