07 Angio Origin

8/10/2019 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


EdmontosaurusEdmontosaurus

Demise of the non-avianDemise of the non-avian

dinosaur lineagedinosaur lineage


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


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


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


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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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


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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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


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- compoundcompound


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)


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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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


morphology phylogeneticmorphology phylogenetic

[some!] analyses place it before[some!] analyses place it before

AmborellaAmborella


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


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)


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


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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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)


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


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


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

07 Angio Origin

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