Magnoliophyta
Magnoliophyta - Angiosperms
- Angiosperms
Survey of Angiosperms
Survey of Angiosperms
—
—
using APG system
using APG system
Amborella trichopoda
Amborella trichopoda
Magnoliophyta
Magnoliophyta - Angiosperms
- Angiosperms
Survey of Angiosperms
Survey of Angiosperms
—
—
using APG system
using APG system
What is the APG classification?
What is the APG classification?
‘
‘
Basal angiosperms
Basal angiosperms
’
’
•
•
ANITA (
ANITA (
‘
‘
basal families
basal families
’
’
)
)
•
•
magnoliid
magnoliid
complex
complex
•
•
monocots
monocots
Eudicots
Eudicots
or
or
tricolpates
tricolpates
(3 pored pollen)
(3 pored pollen)
•
•
caryophyllids
caryophyllids
•
•
rosids
rosids
•
•
asterids
asterids
We will begin our survey of
We will begin our survey of
angiosperms by examining the
angiosperms by examining the
‘
‘basal angiospermsbasal angiosperms’’ - those groups - those groups
that are now shown to be the oldest
that are now shown to be the oldest
diverging
diverging ––paraphyleticparaphyletic!!
We will look at the
We will look at the monocotsmonocots - a - a ‘
‘basal angiosperm groupbasal angiosperm group’’ - at the - at the end of the class
end of the class
Basal Angiosperms
Basal Angiosperms
These include all those shown These include all those shown here
here exceptexcept the the eudicotseudicots whichwhich are the bulk of dicots
are the bulk of dicots
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
polypetaly hypogyny actinomorphic CharlesCharles BesseyBessey’’s orders order RanalesRanales
are most of the
are most of the dicot dicot basal basal angiosperms
angiosperms Magnolia
Magnolia = primitive = primitive
and
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Cronquist
Cronquist’’s subclasss subclass Magnoliidae Magnoliidae . . . .
Cronquist Cronquist
and
and monocotsmonocots
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Exhibit a suite of primitive Exhibit a suite of primitive character states
character states
1. Many parts at each whorl
1. Many parts at each whorl
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Exhibit a suite of primitive Exhibit a suite of primitive character states
character states
1. Many parts at each whorl 1. Many parts at each whorl 2. Separate, unsealed
2. Separate, unsealed carpelscarpels
Drimys
Drimys WinteraceaeWinteraceae
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Exhibit a suite of primitive Exhibit a suite of primitive character states
character states
1. Many parts at each whorl 1. Many parts at each whorl 2. Separate, unsealed carpels 2. Separate, unsealed carpels 3. Follicle fruits
3. Follicle fruits
Leaf-like follicles
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Exhibit a suite of primitive Exhibit a suite of primitive character states
character states
1. Many parts at each whorl 1. Many parts at each whorl 2. Separate, unsealed carpels 2. Separate, unsealed carpels 3. Follicle fruits 3. Follicle fruits Laminar stamens in Laminar stamens in yellow waterlily yellow waterlily 4. Laminar stamens 4. Laminar stamens
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Exhibit a suite of primitive Exhibit a suite of primitive character states
character states
1. Many parts at each whorl 1. Many parts at each whorl 2. Separate, unsealed carpels 2. Separate, unsealed carpels 3. Follicle fruits
3. Follicle fruits 4. Laminar stamens 4. Laminar stamens
5. Tracheids, no vessel elements
5. Tracheids, no vessel elements
What are basal angiosperms?
What are basal angiosperms?
Basal Angiosperms
Basal Angiosperms
Exhibit a suite of primitive Exhibit a suite of primitive character states
character states
1. Many parts at each whorl 1. Many parts at each whorl 2. Separate, unsealed carpels 2. Separate, unsealed carpels 3. Follicle fruits
3. Follicle fruits 4. Laminar stamens 4. Laminar stamens
5. Tracheids, no vessel elements 5. Tracheids, no vessel elements 6. Pollen grains single pored,
6. Pollen grains single pored,
apertured, furrowed;
apertured, furrowed; notnot triaperturate, tricolpate triaperturate, tricolpate
Lilium Lilium (monocot) (monocot)
uniaperturate
uniaperturate The
The ‘‘ANITAANITA’’ group forms a basal grade (paraphyletic) group forms a basal grade (paraphyletic)
‘
‘
ANITA
ANITA
’
’
Basal Angiosperms
Basal Angiosperms
Name derived from initials of members Name derived from initials of members
*Amborellaceae (order Amborellales)
*Amborellaceae (order Amborellales)
‘
‘
A
A
NITA
NITA
’
’
Basal Angiosperms
Basal Angiosperms
Amborella trichopoda
Amborella trichopoda —— only 1 species from New Caledonia only 1 species from New Caledonia
*Amborellaceae (order Amborellales)
*Amborellaceae (order Amborellales)
‘
‘
A
A
NITA
NITA
’
’
Basal Angiosperms
Basal Angiosperms
•
• dioecious (unisexual) evergreen dioecious (unisexual) evergreen shrub with no vessels
shrub with no vessels •
• P 5-8 A P 5-8 A
∞
∞
G G 5-6 5-6 •• perianthperianth spiralledspiralled •
• apocarpicapocarpic, not sealed, not sealed •
• fruits 1 seeded, drupe-like fruits 1 seeded, drupe-like
Illiciaceae
Illiciaceae + + SchisandraceaeSchisandraceae (now combined into latter family) (now combined into latter family)
‘
‘
AN
AN
I
I
TA
TA
’
’
Basal Angiosperms
Basal Angiosperms
•
• aromatic shrubs/vines, used in anise liquers aromatic shrubs/vines, used in anise liquers •
• Asian tropics and subtropics & disjunct in Asian tropics and subtropics & disjunct in Eastern North America
Eastern North America
Illicium
Illicium SchisandraceaeSchisandraceae
Illiciaceae + Schisandraceae
Illiciaceae + Schisandraceae
‘
‘
AN
AN
I
I
TA
TA
’
’
Basal Angiosperms
Basal Angiosperms
P
P
∞
∞
A A∞
∞
GG∞
∞
•
• not sealed carpels, 1 seeded not sealed carpels, 1 seeded follicles
follicles
Trimeniaceae:
Trimeniaceae: Trimenia with 5 spp. (Australasian trees,Trimenia with 5 spp. (Australasian trees,
shrubs, vines)
shrubs, vines)
‘
‘
ANI
ANI
T
T
A
A
’
’
Basal Angiosperms
Basal Angiosperms
P
P
∞
∞
A A∞
∞
GG 1 1 •• filamentous stamens; berry filamentous stamens; berry with one ovule
with one ovule
Austrobaileyaceae: 1 species of tropical Australian
Austrobaileyaceae: 1 species of tropical Australian
evergreen lianaa
evergreen lianaa
‘
‘
ANIT
ANIT
A
A
’
’
Basal Angiosperms
Basal Angiosperms
P
P
∞
∞
A A∞
∞
GG∞
∞
•
• laminar stamens; fly pollinated laminar stamens; fly pollinated
Biogeographical distributions of the A ITA members of
Biogeographical distributions of the A ITA members of
A
ANITA support old notion that NITA support old notion that tropical Australasiatropical Australasia was was centercenter of origin
of origin of extant angiosperms of extant angiosperms
‘
‘
ANITA
ANITA
’
’
Basal Angiosperms
Basal Angiosperms
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
•
• worldwide worldwide
distribution except arid distribution except arid regions
regions •
• little biogeographic little biogeographic structure
structure Distribution of Nymphaeaceae - water lilies
Distribution of Nymphaeaceae - water lilies
•
• water lilies and water lilies and relatives relatives
•
• floating or submersed floating or submersed leaves
leaves •
• air cavities in tissue air cavities in tissue •
• mucilaginous mucilaginous coverings coverings •
• lack of vessels lack of vessels
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
8 genera with specialized ecological aquatic niche
8 genera with specialized ecological aquatic niche
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
Obvious
Obvious ecological convergenceecological convergence in floating aquatics is the rule! in floating aquatics is the rule! Check out live aquatic plant display in Birge lobby.
Check out live aquatic plant display in Birge lobby. Nuphar variegatum Nuphar variegatum - -yellow waterlily yellow waterlily = basal angiosperm = basal angiosperm Nymphoides peltata Nymphoides peltata - -water gentian water gentian = asterid dicot = asterid dicot
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
Order
Order NymphaealesNymphaeales onceonce included all these 3 genera included all these 3 genera Nuphar
Nuphar NelumboNelumbo
Ceratophyllum Ceratophyllum all are unrelated!
all are unrelated!
= ANITA
= ANITA = eudicot= eudicot
= basal angiosperm
= basal angiosperm
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
Nymphaea odorata
Nymphaea odorata - water lily - water lily
CA
4-CA 4-∞ ∞ CO CO∞∞ A A ∞ ∞ GG ( (∞∞)) •• showy flowers with strong showy flowers with strong scent attracting animals scent attracting animals •
• many parts at each whorl many parts at each whorl •
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
•
• showy flowers with strong showy flowers with strong scent attracting animals scent attracting animals •
• many parts at each whorl many parts at each whorl •
• laminar stamens laminar stamens •
• superior, syncarpic pistil superior, syncarpic pistil CA
4-CA 4-∞ ∞ CO CO∞∞ A A ∞ ∞ GG ( (∞∞))
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
*Nymphaeaceae: 2nd lineage after
*Nymphaeaceae: 2nd lineage after Amborella to divergeAmborella to diverge
Nuphar variegata Nuphar variegata -
-yellow pond lily
yellow pond lily •• petaloid sepals, petals reduced petaloid sepals, petals reduced
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
Amazonian
Amazonian VictoriaVictoria with peltate leaves with peltate leaves
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
Cabombaceae - 2 genera often placed in Nymphaeaceae
Cabombaceae - 2 genera often placed in Nymphaeaceae
Brasenia shreberi
Brasenia shreberi - water shield - water shield •••• small clonal floating aquatic small clonal floating aquatic peltate leaves peltate leaves
•
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
Cabombaceae - 2 genera often placed in Nymphaeaceae
Cabombaceae - 2 genera often placed in Nymphaeaceae
Cabomba
Cabomba - fanwort - fanwort •••• submersed and floating leaved submersed and floating leaved dimorphic leaves dimorphic leaves
•
• insect pollinated insect pollinated
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
Molecular phylogeny - secondary increase in floral structures
Molecular phylogeny - secondary increase in floral structures
Don Les & al. 1999
‘
‘
A
A
N
N
ITA
ITA
’
’
Basal Angiosperms
Basal Angiosperms
Hydatellaceae - the enigmatic aquatic finds a home!
Hydatellaceae - the enigmatic aquatic finds a home!
Hydatella
Hydatella TrithuriaTrithuria
•
• 2 Australasian genera of reduced aquatics 2 Australasian genera of reduced aquatics
•
• DNA = monocot near grasses DNA = monocot near grasses
•
• new DNA & morphology = new DNA & morphology = sister to watersister to water lilies
lilies
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Remainder of basal angiosperms
Remainder of basal angiosperms
(except monocots & hornwort) (except monocots & hornwort) •
•
• tropical trees or paleoherbs (cordate leaves) tropical trees or paleoherbs (cordate leaves) •
• aromatic - ethereal oils aromatic - ethereal oils
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Remainder of basal angiosperms
Remainder of basal angiosperms (except monocots & hornwort)(except monocots & hornwort) •
• tropical trees or paleoherbs (cordate leaves) tropical trees or paleoherbs (cordate leaves) •
• aromatic - ethereal oils (“ aromatic - ethereal oils (“ranalianranalian”” smell) smell) •
• beetle or fly pollination common beetle or fly pollination common •
• tropical trees or paleoherbs (cordate leaves) tropical trees or paleoherbs (cordate leaves)
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Magnoliaceae (Magnoliales)
*Magnoliaceae (Magnoliales)
•
• tropical or warm temperate trees or shrubs with tropical or warm temperate trees or shrubs with large, pinnate netted, stipulate leaves
large, pinnate netted, stipulate leaves •
• flowers large and solitary flowers large and solitary
Magnolia grandiflora Magnolia grandiflora
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Magnoliaceae (Magnoliales)
*Magnoliaceae (Magnoliales)
•
• pperianth = tepals spirally arrangederianth = tepals spirally arranged P P ∞∞ A A ∞∞ GG ∞∞ p p p p p p •
• aandroecium of laminar stamensndroecium of laminar stamens
a a a a a a •
• ggynoecium of many separate pistils or carpelsynoecium of many separate pistils or carpels
g g g g g g
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Magnoliaceae (Magnoliales)
*Magnoliaceae (Magnoliales)
•
•fruits of one flower = fruits of one flower = ‘‘conecone’’ or or ‘‘aggregateaggregate’’ of of folliclesfollicles
•
• dehiscent along one suture, derived from one carpel (leaf) dehiscent along one suture, derived from one carpel (leaf) P
P ∞∞ A A ∞∞ GG ∞∞
Magnolia grandiflora Magnolia grandiflora
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Magnoliaceae (Magnoliales)
*Magnoliaceae (Magnoliales)
•
• fruits of one flower = cone of follicles fruits of one flower = cone of follicles
•
• dehiscent along one suture, derived from one carpel (leaf) dehiscent along one suture, derived from one carpel (leaf) P
P ∞∞ A A ∞∞ GG ∞∞
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Magnoliaceae (Magnoliales)
*Magnoliaceae (Magnoliales)
Liriodendron
Liriodendron - tulip tree, yellow - tulip tree, yellow poplar -
poplar - samarasamara fruits fruits
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Annonaceae (Magnoliales) - custard apples
*Annonaceae (Magnoliales) - custard apples
•
• large, woody pantropical family large, woody pantropical family •
• perianth in 3 sets of three tepals perianth in 3 sets of three tepals
P
P 3+3+33+3+3 A A ∞ ∞ GG ∞∞
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Annonaceae (Magnoliales) - custard apples
*Annonaceae (Magnoliales) - custard apples
•
• large, woody pantropical family large, woody pantropical family •
• perianth in 3 sets of three tepals perianth in 3 sets of three tepals •
• fruits aromatic, aggregates of one fruits aromatic, aggregates of one carpeled berries carpeled berries P P 3+3+33+3+3 A A ∞ ∞ GG ∞∞ Annona anon, cherimoya, anon, cherimoya, custard-apple custard-apple
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Annonaceae (Magnoliales) - custard apples
*Annonaceae (Magnoliales) - custard apples
•
• Asimina trilobaAsimina triloba: paw-paw: paw-paw •
• native to eastern North America native to eastern North America •
• flowers fly pollinated; fruits banana-like flowers fly pollinated; fruits banana-like P
P 3+3+33+3+3 A A ∞ ∞ GG ∞∞
Ohio paw-paw fair
Ohio paw-paw fair
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Myristicaceae (Magnoliales) - nutmeg
Myristicaceae (Magnoliales) - nutmeg
•
• Myristica fragransMyristica fragrans - nutmeg, mace - nutmeg, mace
•
• mace from aril mace from aril
•
•nutmeg fromnutmeg from
seed
seed
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Lauraceae (Laurales) - cinnamon, laurel
Lauraceae (Laurales) - cinnamon, laurel
•
• aromatic trees or shrubs aromatic trees or shrubs •
• 3 merous flowers 3 merous flowers
P
P 3+33+3 A 3- A 3-∞∞ GG 11
cinnamon
cinnamon
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Lauraceae (Laurales) - cinnamon, laurel
Lauraceae (Laurales) - cinnamon, laurel
avocado
avocado
P
P 3+33+3 A 3- A 3-∞∞ GG 11 •
• stamens with basal nectar appendages stamens with basal nectar appendages
and
and valved anthersvalved anthers
valved anthers valved anthers
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Lauraceae (Laurales) - cinnamon, laurel
Lauraceae (Laurales) - cinnamon, laurel
avocado
avocado
P
P 3+33+3 A 3- A 3-∞∞ GG 11 •
• stamens with basal nectar appendages stamens with basal nectar appendages
and
and valved anthersvalved anthers
•
• fruit 1 seeded berry or drupe fruit 1 seeded berry or drupe
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Lauraceae (Laurales) - cinnamon, laurel
Lauraceae (Laurales) - cinnamon, laurel
Lindera benzoin
Lindera benzoin - spicebush - spicebush Eastern North America
Eastern North America Laurus nobilis
Laurus nobilis - laurel - laurel
Mediterranean Europe
Mediterranean Europe
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Piperaceae (Piperales) - pepper, pepperomia
Piperaceae (Piperales) - pepper, pepperomia
•
• herbs, vines, shrubs, epiphytes, herbs, vines, shrubs, epiphytes, cordate leaves, bi- or unisexual cordate leaves, bi- or unisexual
P 0 A 3+3 P 0 A 3+3 GG 1 1 Peperomia
Peperomia Piper
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Piperaceae (Piperales) - pepper, pepperomia
Piperaceae (Piperales) - pepper, pepperomia
Piper nigrum -
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Aristolochiaceae (Piperales) - birthwort, wild ginger
*Aristolochiaceae (Piperales) - birthwort, wild ginger
•
• climbing or rhizomatous herbs, cordate leaves climbing or rhizomatous herbs, cordate leaves •
• aromatic, medicinal compounds [“ aromatic, medicinal compounds [“well bornwell born”” family] family] Aristolochia clematis
Aristolochia clematis
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Aristolochiaceae (Piperales) - birthwort, wild ginger
*Aristolochiaceae (Piperales) - birthwort, wild ginger
CA (3) CO 0 A
6-CA (3) CO 0 A 6-∞∞ G (4-6) G (4-6) •
• calyx corolloid calyx corolloid •
• petals absent petals absent •
• fly pollination fly pollination •
• inferior, syncarpic inferior, syncarpic
A. hirta A. hirta
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Aristolochiaceae (Piperales) - birthwort, wild ginger
*Aristolochiaceae (Piperales) - birthwort, wild ginger
•
•Asarum canadensisAsarum canadensis -
-wild ginger
wild ginger
•
• creeping rhizome,creeping rhizome, paired leaves, flowers paired leaves, flowers basal
basal •
• rhizome makes rhizome makes candied ginger candied ginger •
• North American North American Indians used for Indians used for contraceptive contraceptive
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
*Aristolochiaceae (Piperales) - birthwort, wild ginger
*Aristolochiaceae (Piperales) - birthwort, wild ginger
• • 3 petals3 petals reduced to reduced to scales scales •
• seeds with seeds with arils, dispersed arils, dispersed by ants by ants CA (3) CO 0 A 6-CA (3) CO 0 A 6-∞∞ G (4-6) G (4-6)
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Saururaceae (Piperales) - lizard
Saururaceae (Piperales) - lizard’’s-tails-tail P 0 A 3-8 P 0 A 3-8 GG (3-5) (3-5) •
• perennial herbs of north temperate with perennial herbs of north temperate with cordate leaves
cordate leaves •
• reduced flowers on showy inflorescence reduced flowers on showy inflorescence Saururus
Saururus HouttyniaHouttynia
showy bract of showy bract of inflorescence inflorescence Ceratophyllum demersum Ceratophyllum demersum - -hornwort, coon
hornwort, coon’’s-tails-tail
Magnoliid Basal Angiosperms
Magnoliid Basal Angiosperms
Ceratophyllaceae (Ceratophyllales) - hornwort
Ceratophyllaceae (Ceratophyllales) - hornwort
P (8-12) A 5- P (8-12) A 5- ∞∞ GG 1 1 •
• aquatic basal angiosperm near aquatic basal angiosperm near eudicots?
eudicots? •