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    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants

    Angiosperms focus of the courseAngiosperms focus of the course vast majority of plant diversityvast majority of plant diversity

    systematics of other plants examined latersystematics of other plants examined later

    What are the non-angiosperm land plants?What are the non-angiosperm land plants?

    Arthur CronquistArthur Cronquists phyla (= divisions)s phyla (= divisions)

    outdated!outdated!

    most phyla are now called into questionmost phyla are now called into question

    See first pages of Chpts 1 & 3 for moreSee first pages of Chpts 1 & 3 for more

    detail (detail (Plant SystematicsPlant Systematics))

    Fungi?Fungi?

    Fungi collectively are not a natural groupFungi collectively are not a natural group More closely related to animals than to plantsMore closely related to animals than to plants

    Charales - stonewortsCharales - stoneworts

    Green algal lineageGreen algal lineage

    Closest relatives to land plants?Closest relatives to land plants?

    Bryophyta - bryophytesBryophyta - bryophytes

    earliest land plants - non vascularearliest land plants - non vascular

    gametophyte dominant, 16000 speciesgametophyte dominant, 16000 species

    3 lineages3 lineages are they a natural group?are they a natural group?

    hornwortshornworts mossesmosses mossesmosses

    liverwortsliverworts liverwortsliverworts

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    Lycopodiophyta - club mossesLycopodiophyta - club mosses

    3 families, 1150 species3 families, 1150 species

    sporophyte dominant, vascular, free sporingsporophyte dominant, vascular, free sporing are they sister to all other tracheophytes?are they sister to all other tracheophytes?

    IsoetesIsoetes - quillworts- quillworts SelaginellaSelaginella - spike mosses- spike mosses

    LycopodiumLycopodium --

    club mossesclub mosses

    Psilophyta - whisk fernsPsilophyta - whisk ferns

    6 species in 2 genera6 species in 2 genera

    vascular plants, leafless green stemmed,vascular plants, leafless green stemmed,lateral sporangialateral sporangia

    are they primitive vascular plants? or reducedare they primitive vascular plants? or reduced

    ferns?ferns?

    PsilotumPsilotum habithabit PsilotumPsilotum branchbranch PsilotumPsilotum sporangiasporangia

    Equisetophyta - horsetailsEquisetophyta - horsetails

    15 species in15 species inEquisetumEquisetum

    vascular plants, reduced leaves, terminalvascular plants, reduced leaves, terminal

    sporangiasporangia

    what are they related to?what are they related to?

    Polypodiophyta - fernsPolypodiophyta - ferns

    11000 species11000 species

    immense variation in habit and habitatimmense variation in habit and habitat

    spores produced in specialized sporangiaspores produced in specialized sporangia

    need a lot of systematic work - tropics!need a lot of systematic work - tropics!

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    Pinophyta - gymnospermsPinophyta - gymnosperms

    conifersconifers

    870 species870 species

    seed plants but seeds nakedseed plants but seeds naked often divided into 4 phylaoften divided into 4 phyla

    is one closer to angiosperms?is one closer to angiosperms?

    juniperjuniperfraser firfraser firpinepine

    cycadscycads

    Pinophyta - gymnospermsPinophyta - gymnosperms

    870 species870 species

    seed plants but seeds nakedseed plants but seeds naked often divided into 4 phylaoften divided into 4 phyla

    is one closer to angiosperms?is one closer to angiosperms?

    female strobilifemale strobilimale strobilusmale strobilus

    ginkgoginkgo

    Pinophyta - gymnospermsPinophyta - gymnosperms

    870 species870 species

    seed plants but seeds nakedseed plants but seeds naked

    often divided into 4 phylaoften divided into 4 phyla

    is one closer to angiosperms?is one closer to angiosperms?

    ovulesovules male strobilimale strobili

    mature seedsmature seeds

    gnetophytesgnetophytes

    Pinophyta - gymnospermsPinophyta - gymnosperms

    870 species870 species

    seed plants but seeds nakedseed plants but seeds naked

    often divided into 4 phylaoften divided into 4 phyla

    is one closer to angiosperms?is one closer to angiosperms?

    Welwitschia mirabilisWelwitschia mirabilis

    EphedraEphedra GnetumGnetum

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    275,000 + species275,000 + species

    seed plants with seeds encased inseed plants with seeds encased in

    ovaryovary

    flowers aflowers a key innovationkey innovation

    Magnoliophyta - angiospermsMagnoliophyta - angiosperms Angiosperms - Flowering PlantsAngiosperms - Flowering Plants tremendous adaptive radiation ontremendous adaptive radiation on

    land (and back into water)land (and back into water)

    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants tremendous adaptive radiation ontremendous adaptive radiation on

    land (and back into water)land (and back into water)

    Eucalyptus regnansEucalyptus regnans

    (Myrtaceae) over 100m tall and(Myrtaceae) over 100m tall and

    19m dbh19m dbh

    Eucalyptus regnansEucalyptus regnans

    (Myrtaceae) -Tasmania(Myrtaceae) -Tasmania

    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants tremendous adaptive radiation ontremendous adaptive radiation on

    land (and back into water)land (and back into water)

    Wolffia microscopicaWolffia microscopica

    (Araceae) less than 1mm long(Araceae) less than 1mm long

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    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants tremendous adaptive radiation ontremendous adaptive radiation on

    land (and back into water)land (and back into water)Rafflesia arnoldiiRafflesia arnoldii (Rafflesiaceae)(Rafflesiaceae)

    from New Guinea, up to 1m acrossfrom New Guinea, up to 1m across

    and 20lbsand 20lbs

    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants tremendous adaptive radiation ontremendous adaptive radiation on

    land (and back into water)land (and back into water) largest inflorescencelargest inflorescence

    Amorphophallus titanumAmorphophallus titanum (Araceae)(Araceae)

    Who is this famous DutchWho is this famous Dutch

    biologist from the early 20thbiologist from the early 20th

    century who is standing next to thecentury who is standing next to the

    titan arum? . . . and what is hetitan arum? . . . and what is he

    famous for?famous for?

    largest inflorescencelargest inflorescence

    Amorphophallus titanumAmorphophallus titanum (Araceae)(Araceae)

    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants Angiosperms - Flowering PlantsAngiosperms - Flowering Plants tremendous adaptive radiation ontremendous adaptive radiation on

    land (and back into water)land (and back into water)

    largest numbers of flowerslargest numbers of flowers

    CoryphaCorypha palmpalm

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    Magnoliophyta classificationMagnoliophyta classification

    typically divided into twotypically divided into two

    classesclasses dicotsdicots andand monocotsmonocots

    Angiosperms - Flowering PlantsAngiosperms - Flowering Plants

    artificial!artificial!

    we will usewe will use AngiospermAngiosperm

    Phylogeny GroupPhylogeny Group ((APG IIAPG II))

    classification system of 2003classification system of 2003

    The FlowerThe Flower Why Important?Why Important?

    1.1. unlike anything else in other plants & extremelyunlike anything else in other plants & extremely

    variable & co-evolved with animalsvariable & co-evolved with animals

    2.2. floral features used in describing and idfloral features used in describing and idinging

    3.3. plant specimens (herbarium) must include flowers orplant specimens (herbarium) must include flowers or

    derived featuresderived features

    4.4. classification of angiosperms relies on flowersclassification of angiosperms relies on flowers

    The Flower:The Flower: most significant feature of angiospermsmost significant feature of angiosperms

    CalochortusCalochortus - fairy lanterns & mariposas (images: T. Givnish)- fairy lanterns & mariposas (images: T. Givnish)

    The FlowerThe Flower What is it?What is it?

    specialized shootspecialized shoot = stem += stem +

    leaves (folia)leaves (folia)

    shoot is highlyshoot is highly modifiedmodified andand

    determinatedeterminate (ceased to grow)(ceased to grow)

    The FlowerThe Flower What is it?What is it?

    from Schleiden 1855from Schleiden 1855

    foliar theoryfoliar theory of flower - J.W.of flower - J.W.

    von Goethe invon Goethe in Attempt to InterpretAttempt to Interpret

    the Metamorphosis of Plantsthe Metamorphosis of Plants

    (1790)(1790)

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    The FlowerThe Flower What is it?What is it?

    developmental/evolutionary origin of thedevelopmental/evolutionary origin of the

    flower still debatedflower still debated

    2.2. PseudanthialPseudanthial theory -theory -

    compound shoot or conecompound shoot or cone

    anthersanthers

    ovulesovules

    1.1. EuanthialEuanthial theory - (foliartheory - (foliar

    theory) - single shoottheory) - single shoot

    thus, a flower is a specialized shoot that:thus, a flower is a specialized shoot that:

    1.1. isis determinatedeterminate (vs. indeterminate)(vs. indeterminate)

    2.2. has ahas a modified stemmodified stem with compressed internodeswith compressed internodes

    3.3. possessespossesses modified leavesmodified leaves with various functions,with various functions,

    these determined by gene arrays (e.g., ABC model)these determined by gene arrays (e.g., ABC model)

    4.4. often clustered in anoften clustered in an inflorescenceinflorescence (larger branch)(larger branch)

    The FlowerThe Flower What is it?What is it?

    The FlowerThe Flower

    1.1. PedunclePeduncle: floral stalk, the stem: floral stalk, the stem

    supporting the flower; sometimessupporting the flower; sometimes

    referred to as thereferred to as the pedicelpedicel

    read chpt 9 inread chpt 9 in Plant SystematicsPlant Systematics!!

    1st half deals with1st half deals with

    vegetative features - we willvegetative features - we will

    cover in lab 2 next weekcover in lab 2 next week

    2nd half deals with2nd half deals withflowers/fruits - lab 3flowers/fruits - lab 3

    2.2. ReceptacleReceptacle: modified floral: modified floral

    stem or axis from which arisestem or axis from which arise

    the floral appendages orthe floral appendages or

    modified leavesmodified leaves

    The FlowerThe Flower

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    3.3. SepalSepal: the outer whorl of leaves,: the outer whorl of leaves,

    green and protective; collectivelygreen and protective; collectively

    called thecalled the calyxcalyx

    The FlowerThe Flower

    4.4. PetalPetal: the second whorl of: the second whorl of

    leaves, typically brightlyleaves, typically brightly

    colored, attracting pollinators;colored, attracting pollinators;

    collectively called thecollectively called the corollacorolla

    The FlowerThe Flower

    5.5. PerianthPerianth: collective term for: collective term for

    sepals and petalssepals and petals

    TepalsTepals if both similarif both similar oror if onlyif only

    one reduced set (sepals)one reduced set (sepals)

    The FlowerThe Flower

    8.8. StamenStamen: the male structure of: the male structure of

    flower comprisingflower comprising filamentfilament andand

    antheranther

    collectively, stamens are thecollectively, stamens are the

    androeciumandroecium (=(= house ofhouse of

    malesmales))

    can be leaf-like in primitivecan be leaf-like in primitive

    angiospermsangiosperms

    The FlowerThe Flower

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    6.6. FilamentFilament: slender stalk of: slender stalk of

    the stamen supporting thethe stamen supporting the

    anther; permits exsertion ofanther; permits exsertion of

    anther with pollen out ofanther with pollen out of

    flowerflower

    The FlowerThe Flower

    7.7. AntherAnther: fertile portion of: fertile portion of

    stamen that dehisces to releasestamen that dehisces to release

    pollen grainspollen grains; composed of; composed of

    anther sacsanther sacs

    The FlowerThe Flower

    NectariesNectaries often near base ofoften near base of

    stamensstamens

    produceproduce nectar rewardnectar reward forfor

    visitors who will move pollenvisitors who will move pollen

    ((pollinatorspollinators))

    e.g., grass-of-parnassus &e.g., grass-of-parnassus &

    fritillaryfritillary

    The FlowerThe Flower

    13.13. PistilPistil: flask-shaped, female: flask-shaped, female

    structure comprising threestructure comprising three

    main partsmain parts

    often referred to asoften referred to as carpelcarpel(s)(s)

    all pistils (1 or more) areall pistils (1 or more) are

    referred to as thereferred to as the gynoeciumgynoecium

    (=(= house of femaleshouse of females))

    The FlowerThe Flower

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    9.9. OvaryOvary: basal portion of pistil: basal portion of pistil

    that contains ovules; at maturitythat contains ovules; at maturity

    becomesbecomes fruitfruit with seedswith seeds

    10.10. OvulesOvules: fertile portions of: fertile portions of

    pistil that contain a femalepistil that contain a female

    gametophyte (embryo sac);gametophyte (embryo sac);

    develop intodevelop into seedsseeds afterafter

    fertilizationfertilization

    The FlowerThe Flower

    11.11. StyleStyle: slender stalk of pistil: slender stalk of pistil

    above ovary that the pollenabove ovary that the pollen

    tubes must pass through totubes must pass through to

    reach eggs in ovulesreach eggs in ovules

    12.12. StigmaStigma: receptive portion: receptive portion

    at top of style that receivesat top of style that receivesand recognizes pollenand recognizes pollen

    The FlowerThe Flower

    Pollination biologyPollination biology Study of the pollen, its transfer, and movementStudy of the pollen, its transfer, and movementdown the styledown the style

    The FlowerThe Flower

    PistilPistil vs.vs. carpelcarpel

    How do you know?How do you know?

    3 examples3 examples

    Carpels not fusedCarpels not fused

    1.1. MonocarpicMonocarpic

    2.2. ApocarpicApocarpic

    Carpels fusedCarpels fused

    3.3. SyncarpicSyncarpic

    The FlowerThe Flower

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    1 floral1 floral leafleaf in gynoeciumin gynoecium

    This gynoecium isThis gynoecium is

    monocarpicmonocarpic

    (one carpel)(one carpel)

    FoldedFolded leafleaf 1 carpel = 1 pistil1 carpel = 1 pistil

    MonocarpicMonocarpicThe FlowerThe Flower

    This gynoecium isThis gynoecium is

    monocarpicmonocarpic

    (one carpel)(one carpel)

    1 carpel = 1 pistil1 carpel = 1 pistil

    The FlowerThe Flower

    legumeslegumes

    If 9If 9 leavesleaves in one flowerin one flower

    eacheach separatelyseparately formsforms

    carpels,carpels,

    then the flower has 9then the flower has 9

    carpels and 9 pistils,carpels and 9 pistils,

    gynoecium isgynoecium is apocarpicapocarpic

    (separate carpels)(separate carpels)

    Caltha palustris -Caltha palustris - Marsh marigoldMarsh marigold

    9 fruits (pistils) from 1 flower9 fruits (pistils) from 1 flowerGynoecium isGynoecium is apocarpicapocarpic with 9 carpels or 9 pistilswith 9 carpels or 9 pistils

    ApocarpicApocarpic

    The FlowerThe Flower

    3 floral3 floral leavesleaves inin

    gynoeciumgynoecium fusefuse

    This gynoecium isThis gynoecium is

    syncarpicsyncarpicThis gynoecium isThis gynoecium is

    syncarpicsyncarpic

    SyncarpicSyncarpic

    The FlowerThe Flower

    3 carpels = 1 pistil3 carpels = 1 pistil

    3 styles3 styles

    3 carpels = 1 pistil3 carpels = 1 pistil

    1 style1 style

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    Placentation typesPlacentation types- arrangement of ovules,- arrangement of ovules,

    provides hints to the number of carpelsprovides hints to the number of carpels

    MarginalMarginal- found in- found in

    almost all monocarpic oralmost all monocarpic or

    apocarpic pistilsapocarpic pistils

    AxileAxile- found in some- found in some

    syncarpic pistilssyncarpic pistils

    The FlowerThe Flower

    ParietalParietal- found in- found in

    some syncarpicsome syncarpic

    pistilspistils

    The FlowerThe Flower

    Placentation typesPlacentation types- arrangement of ovules,- arrangement of ovules,

    provides hints to the number of carpelsprovides hints to the number of carpels

    ParietalParietal- found in- found in

    some syncarpicsome syncarpic

    pistilspistils

    The FlowerThe Flower

    Placentation typesPlacentation types- arrangement of ovules,- arrangement of ovules,

    provides hints to the number of carpelsprovides hints to the number of carpels

    Free-centralFree-central- found in- found in

    a few syncarpic pistilsa few syncarpic pistils

    ParietalParietal- found in- found in

    some syncarpicsome syncarpic

    pistilspistils

    The FlowerThe Flower

    Placentation typesPlacentation types- arrangement of ovules,- arrangement of ovules,

    provides hints to the number of carpelsprovides hints to the number of carpels

    Free-centralFree-central- found in- found in

    a few syncarpic pistilsa few syncarpic pistils

    BasalBasal- found in- found in

    some monocarpic,some monocarpic,

    apocarpic, orapocarpic, or

    syncarpic pistilssyncarpic pistils

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    perianthperianth spiralledspiralled

    Common in primitiveCommon in primitive

    angiospermsangiosperms

    perianthperianth 5-merous5-merous

    Common in dicotsCommon in dicots

    Numerical planNumerical plan- merosity, arrangement of perianth- merosity, arrangement of perianth

    not necessarily stamens or carpelsnot necessarily stamens or carpels

    The FlowerThe Flower

    perianthperianth 4-merous4-merous

    Occasional in dicotsOccasional in dicots

    perianthperianth 3-merous3-merous

    Common in monocots & someCommon in monocots & some

    primitive angiospermsprimitive angiosperms

    The FlowerThe Flower

    Numerical planNumerical plan- merosity, arrangement of perianth- merosity, arrangement of perianth

    not necessarily stamens or carpelsnot necessarily stamens or carpels

    FlowersFlowers radiallyradially

    symmetricalsymmetrical

    FlowersFlowers actinomorphicactinomorphic

    FlowersFlowers bilaterallybilaterally

    symmetricalsymmetrical

    FlowersFlowers zygomorphiczygomorphic

    Symmetry planSymmetry plan- perianth arrangement important in- perianth arrangement important in

    pollination biologypollination biology

    The FlowerThe Flower

    ConnationConnation: fusion of floral: fusion of floral

    parts from theparts from the same whorlsame whorl

    Fusion of carpelsFusion of carpels Syncarpic pistilSyncarpic pistil

    Fusion of stamensFusion of stamens Staminal tubeStaminal tube Fusion of petalsFusion of petals Corolla tubeCorolla tube

    The FlowerThe Flower

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    AdnationAdnation: fusion of floral: fusion of floral

    parts fromparts from different whorlsdifferent whorls

    The FlowerThe Flower

    SimpleSimple adnationadnation

    Stamens fused onto innerStamens fused onto inner

    surface of fused (connation)surface of fused (connation)

    petalspetals

    ComplexComplex adnationadnation

    Sepals, petals, and stamens fuseSepals, petals, and stamens fuse

    to form ato form a hypanthiumhypanthium

    AdnationAdnation: fusion of floral: fusion of floral

    parts fromparts from different whorlsdifferent whorls

    No adnation!No adnation!

    Connation (fusion of similarConnation (fusion of similar

    parts) may or may not occurparts) may or may not occur

    The FlowerThe Flower

    e.g.,e.g., DrimysDrimys & sandwort& sandwort

    Adnation of calyx,Adnation of calyx,corolla, & stamenscorolla, & stamens

    == hypanthiumhypanthium

    The FlowerThe Flower

    AdnationAdnation: fusion of floral: fusion of floral

    parts fromparts from different whorlsdifferent whorlse.g., cherry & rosee.g., cherry & rose

    The FlowerThe Flower

    AdnationAdnation: fusion of floral: fusion of floral

    parts fromparts from different whorlsdifferent whorlse.g., horse gentian & fuchsiae.g., horse gentian & fuchsia

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    Oenothera biennisOenothera biennis

    Evening primroseEvening primrose

    OnagraceaeOnagraceae

    4 sepals (4 sepals (CACAlyx)lyx)

    4 petals (4 petals (COCOrolla)rolla)

    8 stamens (8 stamens (AAndroecium)ndroecium)

    4 carpels (4 carpels (GGynoecium)ynoecium)

    Floral formulaFloral formula- shorthand notation- shorthand notation

    The FlowerThe Flower

    Oenothera biennisOenothera biennis

    Evening primroseEvening primrose

    OnagraceaeOnagraceae

    4 sepals (4 sepals (CACAlyx)lyx)

    4 petals (4 petals (COCOrolla)rolla)

    8 stamens (8 stamens (AAndroecium)ndroecium)

    4 carpels (4 carpels (GGynoecium)ynoecium)

    Floral formulaFloral formula- shorthand notation- shorthand notation

    The FlowerThe Flower

    Carpels fused = 1 pistilCarpels fused = 1 pistil

    Oenothera biennisOenothera biennis

    Evening primroseEvening primrose

    OnagraceaeOnagraceae

    4 sepals (4 sepals (CACAlyx)lyx)

    4 petals (4 petals (COCOrolla)rolla)

    8 stamens (8 stamens (AAndroecium)ndroecium)

    4 carpels (4 carpels (GGynoecium)ynoecium)

    Floral formulaFloral formula- shorthand notation- shorthand notation

    The FlowerThe Flower

    Carpels fused = 1 pistilCarpels fused = 1 pistil

    Ovary inferiorOvary inferior

    Oenothera biennisOenothera biennis

    Evening primroseEvening primrose

    OnagraceaeOnagraceae

    4 sepals (4 sepals (CACAlyx)lyx)

    4 petals (4 petals (COCOrolla)rolla)

    8 stamens (8 stamens (AAndroecium)ndroecium)

    4 carpels (4 carpels (GGynoecium)ynoecium)

    Floral formulaFloral formula- shorthand notation- shorthand notation

    The FlowerThe Flower

    Carpels fused = 1 pistilCarpels fused = 1 pistil

    Ovary inferiorOvary inferior

    Hypanthium (+ hypanthium tube)Hypanthium (+ hypanthium tube)

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    TheThe flowerflower what is it?what is it?

    a flower is a specialized shoot that:a flower is a specialized shoot that:

    1.1. isis determinatedeterminate (vs. indeterminate)(vs. indeterminate)

    2.2. has ahas a modified stemmodified stem with compressed internodeswith compressed internodes

    3.3. possessespossesses modified leavesmodified leaves with various functions,with various functions,

    these determined by gene arrays (e.g., ABC model)these determined by gene arrays (e.g., ABC model)

    4.4. often clustered in anoften clustered in an inflorescenceinflorescence (larger branch)(larger branch)

    Angiosperms -Angiosperms -

    Floral StructureFloral Structure

    See second half of Chpt 9 for more detailSee second half of Chpt 9 for more detail

    ((Plant SystematicsPlant Systematics))