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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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