07 angio origin
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DarwinDarwins Abominables AbominableMysteryMystery
. . . origin of angiosperms . . .. . . origin of angiosperms . . .
Great Mysteries to ZoologistsGreat Mysteries to Zoologists
ArchaeopteryxArchaeopteryxRise of the birds from a dinosaur lineageRise of the birds from a dinosaur lineage
Great Mysteries to ZoologistsGreat Mysteries to Zoologists
EdmontosaurusEdmontosaurus
Demise of the non-avianDemise of the non-avian
dinosaur lineagedinosaur lineage
Great Mysteries to ZoologistsGreat Mysteries to Zoologists
Eocene inEocene in
KansasKansas
Adaptive radiation ofAdaptive radiation of
mammals after dinosaursmammals after dinosaurs
KT eventKT event
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Greatest Mystery to BotanistsGreatest Mystery to Botanists
Origin and rise ofOrigin and rise of
angiospermsangiosperms
Angiosperm DominanceAngiosperm Dominance
Angiosperm DominanceAngiosperm Dominance Angiosperm DominanceAngiosperm Dominance
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Angiosperm DominanceAngiosperm Dominance Angiosperm DominanceAngiosperm Dominance
Gymnosperm DominanceGymnosperm Dominance Gymnosperm DominanceGymnosperm Dominance
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Angiosperms have NOT always dominatedAngiosperms have NOT always dominated Fossil RecordFossil Recordas seen by Charles Darwinas seen by Charles Darwin
ferns, cycads,ferns, cycads, ginkosginkos, conifers -, conifers -
thethe Triassic/JurassicTriassic/Jurassic
flowering plants suddenly showflowering plants suddenly show
up atup at start of Cretaceousstart of Cretaceous
tree-like lycopods, horsetails,tree-like lycopods, horsetails,primitive gymnosperms - theprimitive gymnosperms - the
CarboniferousCarboniferous
Fossil RecordFossil Record
by mid-Cretaceous, 50 familiesby mid-Cretaceous, 50 families
of angiosperms seenof angiosperms seen
including 5 monocotincluding 5 monocot
including 4 ament/catkinincluding 4 ament/catkinbearingbearing
as seen by Charles Darwinas seen by Charles Darwin
Fossil RecordFossil Record
by mid-Cretaceous, angiosperms also dominate the face ofby mid-Cretaceous, angiosperms also dominate the face of
the earth (based on fossil diversity)the earth (based on fossil diversity)
as seen by Charles Darwinas seen by Charles Darwin
% of total fossils
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the Abominable Mysterythe Abominable Mystery
The rapid development, as far as we canThe rapid development, as far as we can
judge, of all the higher plants within recentjudge, of all the higher plants within recent
geological time isgeological time is an abominable mysteryan abominable mystery
Director of the KewDirector of the KewRoyal Botanic GardenRoyal Botanic Gardenand good friend ofand good friend ofDarwin (the onlyDarwin (the onlyacknowledged person inacknowledged person inthethe Origin of SpeciesOrigin of Species))
Joseph Dalton HookerJoseph Dalton Hooker
(Darwin, 1879, in a letter to(Darwin, 1879, in a letter to
Hooker)Hooker)
the Abominable Mysterythe Abominable Mystery
(page 3, letter of 22 July 1879)(page 3, letter of 22 July 1879)
- Continues with speculations on- Continues with speculations on
how to answer the mysteryhow to answer the mystery
originated in alpine
conditions
arose in response to rise
of flower-frequenting
insects
originated in isolated
tropical island
originated in alpineoriginated in alpine
conditionsconditions
arose in response to risearose in response to rise
ofof flower-frequentingflower-frequenting
insectsinsects
originated in isolatedoriginated in isolated
tropical islandtropical island
the 2012 Questionsthe 2012 Questions
1.1. WhenWhendid the Angiosperms arise?did the Angiosperms arise?
2.2. WhatWhatwere the first Angiosperms?were the first Angiosperms?
3.3. WhereWheredid the Angiosperm arise?did the Angiosperm arise?
4.4. From whatFrom whatGymnosperm cladeGymnosperm clade
did the Angiosperms arise ?did the Angiosperms arise ?
5.5. WhyWhydid they take over the worlddid they take over the worlds flora?s flora?
2009 AJB
volume
2009 AJB2009 AJB
volumevolume
the 2012 Evidencethe 2012 Evidence
1.1. Modern fossil recordModern fossil record
2.2. Geographical distributionsGeographical distributions
3.3. Morphological phylogeneticsMorphological phylogenetics
4.4. Evo-devo studies of flowersEvo-devo studies of flowers
5.5. Molecular phylogeneticsMolecular phylogenetics
6.6. MolecularMolecular clocksclocks
PollenPollen
LeavesLeaves
FlowersFlowers
FruitFruit
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Pollen RecordPollen Record
ubiquitousubiquitous- preserves well due to- preserves well due to exineexinelayerlayer
oftenoften diagnosticdiagnosticto specific gymnosperm or angiospermto specific gymnosperm or angiosperm
groupsgroups
Pollen RecordPollen Record
ubiquitousubiquitous- preserves well due to- preserves well due to exineexinelayerlayer
oftenoften diagnosticdiagnosticto specific gymnosperm or angiospermto specific gymnosperm or angiosperm
groupsgroups
butbut different levelsdifferent levelsof production and fossilizationof production and fossilization
abundant pine pollen in lake sedimentsabundant pine pollen in lake sediments
littlelittle SenecioSeneciopollen in tropicspollen in tropics
Pollen RecordPollen Record
gymnosperm vs. early angiosperm pollen differentiationgymnosperm vs. early angiosperm pollen differentiation
oftenoften requires TEM vs. SEMrequires TEM vs. SEMvisualization - both one poredvisualization - both one pored
Pollen RecordPollen Record
all pre-Cretaceousall pre-Cretaceouspollen = gymnospermpollen = gymnosperm
Barr.-Aptian (Barr.-Aptian (125mya125mya))
= oldest tricolpate pollen= oldest tricolpate pollen
((eudicotseudicots))
Neocomian (Neocomian (130mya130mya))
= oldest angiosperm= oldest angiosperm
single pored pollensingle pored pollen
((basal angiospermsbasal angiosperms))
Jim Doyle - UC DavisJim Doyle - UC Davis
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Pollen RecordPollen Record
magnoliidsmagnoliids
monocotsmonocots
cordate-leavedcordate-leavedeudicotseudicots
aments - windaments - windpollinatedpollinated
Albian (110 mya)Albian (110 mya)
diversitydiversity
Upper Cretaceous (100Upper Cretaceous (100
mya) -mya) - angiospermangiosperm
pollen dominatespollen dominates
Pollen RecordPollen Recordpollen diversification continues through Upper Cretaceouspollen diversification continues through Upper Cretaceous
into Tertiaryinto Tertiary
familyfamily AsteraceaeAsteraceae
first seen in midfirst seen in mid
Eocene: 47myaEocene: 47mya
genusgenus FuchsiaFuchsia
((OnagraceaeOnagraceae) first) first
seen 30mya inseen 30mya in
OligoceneOligocene
Leaf RecordLeaf Recordconsistent trends emerge with leaf fossilsconsistent trends emerge with leaf fossils
Neocomian (Neocomian (130mya130mya))
RogersiaRogersia (basal(basal
angiosperm) simple,angiosperm) simple,
pinnately veined, entirepinnately veined, entire
Leaf RecordLeaf Recordconsistent trends emerge with leaf fossilsconsistent trends emerge with leaf fossils
Aptian to Albian (Aptian to Albian (120-120-
110mya110mya) =) = magnoliidsmagnoliids
(pinnate veins),(pinnate veins), cordatescordates
(palmate veins),(palmate veins),
monocotsmonocots(parallel(parallel
veins)veins)
early Aptian (early Aptian (125mya125mya))
ArchaefructusArchaefructus(basal(basal
angiosperm) palmatelyangiosperm) palmately
compoundcompound
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Leaf RecordLeaf Recordconsistent trends emerge with leaf fossilsconsistent trends emerge with leaf fossils
AptianAptiantoto AlbianAlbian((120-120-
110mya110mya) =) = magnoliidsmagnoliids
(pinnate veins),(pinnate veins), cordatescordates
(palmate veins),(palmate veins),
monocotsmonocots(parallel(parallel
veins)veins)
early Aptian (early Aptian (125mya125mya))
ArchaefructusArchaefructus(basal(basal
angiosperm) palmatelyangiosperm) palmately
compoundcompound
great leaf diversity within 15mygreat leaf diversity within 15my
Leaf RecordLeaf Recordconsistent trends emerge with leaf fossilsconsistent trends emerge with leaf fossils
by Upper Cretaceousby Upper Cretaceous
(100mya) a variety of(100mya) a variety of
primitive eudicot leavesprimitive eudicot leaves
are seenare seen
PlatanoidPlatanoid- lobed- lobed
SapindopsisSapindopsis- compound- compound
Leaf RecordLeaf Recordconsistent trends emerge with leaf fossilsconsistent trends emerge with leaf fossils
by Upper Cretaceousby Upper Cretaceous
(100mya) a variety of(100mya) a variety of
primitive eudicot leavesprimitive eudicot leaves
are seenare seen
Flower RecordFlower Recordthethe Magnolia = primitiveMagnolia = primitiveidea has biased the wayidea has biased the way
paleobotanists have looked at the fossil recordpaleobotanists have looked at the fossil record
1.1. bisexual flowerbisexual flower
spirally arranged stamens &spirally arranged stamens &
carpelscarpels
perianth partsperianth parts
4.4. cone-like receptaclecone-like receptacle
5.5. beetle pollinationbeetle pollination
what does the fossil recordwhat does the fossil record
actually say?actually say?
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Flower RecordFlower Recordlarge flowered, insect-pollinated flowers are seen (such aslarge flowered, insect-pollinated flowers are seen (such as
these 98-90these 98-90 myamyamid-late Cretaceous fossils) . . .mid-late Cretaceous fossils) . . .
ArchaeanthusArchaeanthus((MagnoliaceaeMagnoliaceae))
from Kansas 98-95myafrom Kansas 98-95mya
MagnoliaceaeMagnoliaceae withwith
stingless beestingless bee90mya90mya
Flower RecordFlower Recordlarge flowered, insect-pollinated flowers are seen (such aslarge flowered, insect-pollinated flowers are seen (such as
these 95-85 mya late Cretaceous fossils) . . .these 95-85 mya late Cretaceous fossils) . . . . . .. . . but the earliest and mostbut the earliest and most
numerous are small, bisexual ornumerous are small, bisexual or
unisexual, wind or insect-pollinatedunisexual, wind or insect-pollinated
Flower RecordFlower Recordwhat are the earliest fossil flowers?what are the earliest fossil flowers?
1990 - Leo Hickey1990 - Leo Hickey
120 mya - Australia120 mya - Australia
small, unisexual flowerssmall, unisexual flowers
placed into Piperalesplaced into Piperales
(pepper, wild ginger)(pepper, wild ginger)
Flower RecordFlower Recordwhat are the earliest fossil flowers?what are the earliest fossil flowers?
1996 - Chris Hill1996 - Chris Hill
Bevhalstia pebiaBevhalstia pebia
130 mya - England130 mya - England
small, 25cm aquaticsmall, 25cm aquatic
herbherb
dissected leavesdissected leaves
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Flower RecordFlower Recordwhat are the earliest fossil flowers?what are the earliest fossil flowers?
1998 - David Dilcher & Chinese1998 - David Dilcher & Chinese
colleaguescolleaguesread Sun et al.!read Sun et al.!
ArchaefructusArchaefructus
125 [135 1125 [135 1stst]] myamya- China- China
small, dissected leavessmall, dissected leaves
stamensstamensandand carpelscarpelson long axison long axis
Flower RecordFlower Recordwhat are the earliest fossil flowers?what are the earliest fossil flowers?
morphology phylogeneticmorphology phylogenetic
[some!] analyses place it before[some!] analyses place it before
AmborellaAmborella
Flower RecordFlower Recordwhat are the earliest fossil flowers?what are the earliest fossil flowers?
2011 - David2011 - David DilcherDilcher& Chinese& Chinese
colleagues againcolleagues again
LeefructusLeefructus
122mya - China122mya - China
firstfirst EudicotEudicot
RanunculaceaeRanunculaceae??
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
1. When1. Whendid the Angiosperms arise?did the Angiosperms arise?
Fossils - after boundary of Jurassic and Cretaceous -Fossils - after boundary of Jurassic and Cretaceous -
130mya130mya
DNA - some molecular clocks suggest >200myaDNA - some molecular clocks suggest >200mya
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MagallMagallnn20102010
Stem lineageStem lineage
stem nodestem node
with livingwith living
gymnospermsgymnosperms
crown nodecrown node
of livingof living
angiospermsangiosperms
350mya350mya
225mya225mya
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
1. When1. Whendid the Angiosperms arise?did the Angiosperms arise?
perhaps older but unseen (in fossil record) radiationperhaps older but unseen (in fossil record) radiation
of angiospermsof angiosperms
perhaps older radiation but we canperhaps older radiation but we cant tell them apartt tell them apart
from ancestors (share features of gymnosperms andfrom ancestors (share features of gymnosperms and
some but not all of angiosperms)some but not all of angiosperms)
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
2. What2. What were the first Angiosperms?were the first Angiosperms?
MagnoliaMagnolia= primitive= primitivenot justifiednot justified
AmborellaAmborellaand water lilies are firstand water lilies are first extantextanttaxa totaxa to
separateseparate
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
2. What2. What were the first Angiosperms?were the first Angiosperms?
MagnoliaMagnolia= primitive= primitivenot justifiednot justified
AmborellaAmborellaand water lilies are firstand water lilies are first extantextanttaxa totaxa to
separateseparate
earliestearliest extinctextinctfossils are small, probably aquaticfossils are small, probably aquatic
plantsplants
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Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
3. Where3. Wheredid the Angiosperms arise?did the Angiosperms arise?
Australasia if based on earliest diverging extantAustralasia if based on earliest diverging extant
familiesfamilies
earliest fossils come from many areas - China,earliest fossils come from many areas - China,
England, Australia (most tropical or subtropical orEngland, Australia (most tropical or subtropical or
warm temperate in early Cretaceous)warm temperate in early Cretaceous)
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
3. Where3. Wheredid the Angiosperms arise?did the Angiosperms arise?
likely in wet margins of gymnosperm dominatedlikely in wet margins of gymnosperm dominated
forestsforests
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
4. From4. From what Gymnosperms did they arise?what Gymnosperms did they arise?
no consensus based on extant lineages!no consensus based on extant lineages!
GnetalesGnetaleshave been the favorites for some timehave been the favorites for some time
(vessels, double fertilization, broad leaves) . . .(vessels, double fertilization, broad leaves) . . .
GnetumGnetum
EphedraEphedra
WelwitschiaWelwitschia
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but thisbut this AnthophyteAnthophytehypothesishypothesisis stronglyis strongly
rejectedrejectedby DNA sequence data!by DNA sequence data!
more likely that Angiosperms arose from a nowmore likely that Angiosperms arose from a now
extinct Gymnosperm lineage such asextinct Gymnosperm lineage such as BennettitalesBennettitalesoror
otherother seed fernseed ferngroupsgroups
Summary of AngiospermSummary of Angiosperm
EvolutionEvolution
5. Why5. Why did Angiosperms dominate quickly?did Angiosperms dominate quickly?
BIG story! We will deal with it throughout theBIG story! We will deal with it throughout the
coursecourse
vessel elements?vessel elements?
mycorrhizalmycorrhizalinteractions with fungi?interactions with fungi?
the flower as athe flower as a key innovationkey innovation??
genome duplication(s)?genome duplication(s)?
co-evolution with animal pollinators?co-evolution with animal pollinators?
does thedoes the timingtimingof Angiosperm & pollinator radiations fit?of Angiosperm & pollinator radiations fit?
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does thedoes the timingtimingof Angiosperm & pollinator radiations fit?of Angiosperm & pollinator radiations fit?
No & YesNo & Yes AngiospermsAngiosperms co-co-
opted already diverging animalopted already diverging animal
lineageslineagesin Early Cretaceousin Early Cretaceous
AngiospermAngiosperm
evolutionevolution
PollinatorPollinator
evolutionevolution
DarwinDarwins Abominables Abominable
MysteryMystery
. . would he be satisfied if he. . would he be satisfied if he
had lived tohad lived to be 203be 203 ? . .? . .
Yes . . . but questionsYes . . . but questions
remainremain