Angiosperms - Flowering Plants Fungi?

• Fungi collectively are not a natural group

Angiosperms focus of the course
• More closely related to animals than to plants
• vast majority of plant diversity
• systematics of other plants examined later

What are the non-angiosperm land plants?

• Arthur Cronquist’
Cronquist’s phyla (= divisions)
• outdated!
• most phyla are now called into question

See first pages of Chpts 1 & 3 for more

detail (Plant
(Plant Systematics)
Systematics)

Charales - stoneworts Bryophyta - bryophytes

• earliest land plants - non vascular
• Green algal lineage • gametophyte dominant, 16000 species
• Closest relatives to land plants? • 3 lineages — are they a natural group?
hornworts mosses mosses

liverworts liverworts
 Lycopodiophyta - club mosses Psilophyta - whisk ferns
• 3 families, 1150 species • 6 species in 2 genera
• sporophyte dominant, vascular, free sporing • vascular plants, leafless green stemmed,
• are they sister to all other tracheophytes? lateral sporangia
• are they primitive vascular plants? or reduced
Lycopodium -
club mosses ferns?

Isoetes - quillworts Selaginella - spike mosses

Psilotum habit Psilotum branch Psilotum sporangia

Equisetophyta - horsetails Polypodiophyta - ferns

• 15 species in Equisetum • 11000 species
• vascular plants, reduced leaves, terminal • immense variation in habit and habitat
sporangia • spores produced in specialized sporangia
• what are they related to? • need a lot of systematic work - tropics!
 Pinophyta - gymnosperms Pinophyta - gymnosperms
• 870 species • 870 species
• seed plants but seeds naked • seed plants but seeds naked
conifers
• often divided into 4 phyla • often divided into 4 phyla
• is one closer to angiosperms? • is one closer to angiosperms?

cycads

pine fraser fir juniper male strobilus female strobili

Pinophyta - gymnosperms Pinophyta - gymnosperms

• 870 species • 870 species
• seed plants but seeds naked • seed plants but seeds naked
• often divided into 4 phyla • often divided into 4 phyla
• is one closer to angiosperms? • is one closer to angiosperms?

gnetophytes

mature seeds

ginkgo

Ephedra Gnetum
ovules male strobili
Welwitschia mirabilis
 Magnoliophyta - angiosperms Angiosperms - Flowering Plants
• tremendous adaptive radiation on
land (and back into water)

• 275,000 + species
• seed plants with seeds encased in
ovary
• flowers a “key innovation”
innovation”

Angiosperms - Flowering Plants Angiosperms - Flowering Plants

• tremendous adaptive radiation on • tremendous adaptive radiation on
land (and back into water) land (and back into water)
• Eucalyptus regnans • Wolffia microscopica
(Myrtaceae) over 100m tall and (Araceae) less than 1mm long
19m dbh
Eucalyptus regnans
(Myrtaceae) -Tasmania
 Angiosperms - Flowering Plants Angiosperms - Flowering Plants
• tremendous adaptive radiation on • tremendous adaptive radiation on
land (and back into water) land (and back into water)
• Rafflesia arnoldii (Rafflesiaceae) • largest inflorescence
from New Guinea, up to 1m across Amorphophallus titanum (Araceae)
and 20lbs

Angiosperms - Flowering Plants Angiosperms - Flowering Plants

• tremendous adaptive radiation on
land (and back into water)
• largest inflorescence • largest numbers of flowers —
Amorphophallus titanum (Araceae) Corypha palm

Who is this famous Dutch

biologist from the early 20th
century who is standing next to the
titan arum? . . . and what is he
famous for?
 Angiosperms - Flowering Plants The Flower — Why Important?
The Flower: most significant feature of angiosperms
Magnoliophyta classification 1. unlike anything else in other plants & extremely
variable & co-evolved with animals
• typically divided into two 2. floral features used in describing and id’
id’ing
classes — dicots and monocots 3. plant specimens (herbarium) must include flowers or
derived features
• artificial! 4. classification of angiosperms relies on flowers

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

• we will use Angiosperm
Phylogeny Group (APG(APG II)
II)
classification system of 2003

The Flower — What is it? The Flower — What is it?

• “foliar theory”
theory” of flower - J.W.
• specialized shoot = stem +
von Goethe in “Attempt to Interpret
leaves (folia)
the Metamorphosis of Plants”
Plants”
(1790)
• shoot is highly modified and
determinate (ceased to grow)

from Schleiden 1855

 The Flower — What is it? The Flower — What is it?
• developmental/evolutionary origin of the • thus, a flower is a specialized shoot that:
flower still debated
1. is determinate (vs. indeterminate)
1. Euanthial theory - (foliar
theory) - single shoot
2. has a modified stem with compressed internodes
anthers
ovules 3. possesses modified leaves with various functions,
these determined by gene arrays (e.g., ABC model)
2. Pseudanthial theory -
compound shoot or cone 4. often clustered in an inflorescence (larger branch)

The Flower The Flower

read chpt 9 in Plant Systematics!
Systematics!
• 1st half deals with
vegetative features - we will
cover in lab 2 next week
• 2nd half deals with
flowers/fruits - lab 3

2. Receptacle:
Receptacle: modified floral
1. Peduncle:
Peduncle: floral stalk, the stem stem or axis from which arise
supporting the flower; sometimes the floral appendages or
referred to as the pedicel modified leaves
The Flower The Flower

4. Petal:
Petal: the second whorl of
3. Sepal:
Sepal: the outer whorl of leaves,
leaves, typically brightly
green and protective; collectively
colored, attracting pollinators;
called the calyx
collectively called the corolla

The Flower The Flower

8. Stamen:
Stamen: the male structure of
flower comprising filament and
anther

• collectively, stamens are the

5. Perianth:
Perianth: collective term for androecium (= ‘house of
sepals and petals males’
males’)

Tepals if both similar or if only • can be leaf-like in primitive

one reduced set (sepals) angiosperms
The Flower The Flower

6. Filament:
Filament: slender stalk of 7. Anther:
Anther: fertile portion of
the stamen supporting the stamen that dehisces to release
anther; permits exsertion of pollen grains;
grains; composed of
anther with pollen out of anther sacs
flower

The Flower The Flower

13. Pistil:
Pistil: flask-shaped, female
structure comprising three
main parts
Nectaries often near base of
stamens • often referred to as carpel(s)
carpel(s)

• produce nectar reward for • all pistils (1 or more) are

visitors who will move pollen referred to as the gynoecium
(‘pollinators’
pollinators’) (= ‘house of females’
females’)

e.g., grass-of-parnassus &

fritillary
 The Flower The Flower
12. Stigma:
Stigma: receptive portion
at top of style that receives
and recognizes pollen
11. Style:
Style: slender stalk of pistil
above ovary that the pollen
9. Ovary:
Ovary: basal portion of pistil tubes must pass through to
that contains ovules; at maturity reach eggs in ovules
becomes fruit with seeds

10. Ovules:
Ovules: fertile portions of
pistil that contain a female
gametophyte (embryo sac);
develop into seeds after
fertilization

The Flower The Flower

Study of the pollen, its transfer, and movement
Pollination biology Pistil vs. carpel
down the style

How do you know?

3 examples

Carpels not fused

1. Monocarpic
2. Apocarpic
Carpels fused
3. Syncarpic
 The Flower The Flower
Monocarpic

1 floral ‘leaf’
leaf’ in gynoecium Folded ‘leaf’
leaf’ 1 carpel = 1 pistil 1 carpel = 1 pistil

This gynoecium is This gynoecium is

monocarpic monocarpic
(one carpel) legumes (one carpel)

The Flower The Flower

Apocarpic Syncarpic

• If 9 ‘leaves’
leaves’ in one flower
each separately forms
carpels,

• then the flower has 9

carpels and 9 pistils,

• gynoecium is apocarpic
(separate carpels)

3 floral ‘leaves’
leaves’ in 3 carpels = 1 pistil 3 carpels = 1 pistil
Caltha palustris - Marsh marigold gynoecium fuse 3 styles 1 style
9 fruits (pistils) from 1 flower
Gynoecium is apocarpic with 9 carpels or 9 pistils This gynoecium is This gynoecium is
syncarpic syncarpic
 The Flower The Flower
Placentation types - arrangement of ovules, Placentation types - arrangement of ovules,
provides hints to the number of carpels provides hints to the number of carpels

Marginal - found in Parietal - found in

almost all monocarpic or some syncarpic
apocarpic pistils pistils

Axile - found in some

syncarpic pistils

The Flower The Flower

Placentation types - arrangement of ovules, Placentation types - arrangement of ovules,
provides hints to the number of carpels provides hints to the number of carpels

Parietal - found in Parietal - found in

some syncarpic some syncarpic
pistils pistils

Free-central - found in Free-central - found in

a few syncarpic pistils a few syncarpic pistils

Basal - found in
some monocarpic,
apocarpic, or
syncarpic pistils
 The Flower The Flower
Numerical plan - merosity, arrangement of perianth Numerical plan - merosity, arrangement of perianth
• not necessarily stamens or carpels • not necessarily stamens or carpels

perianth spiralled perianth 5-merous perianth 4-merous perianth 3-merous

Common in primitive Common in dicots Occasional in dicots Common in monocots & some
angiosperms primitive angiosperms

The Flower The Flower

Symmetry plan - perianth arrangement important in Connation:
Connation: fusion of floral
pollination biology parts from the same whorl

Fusion of carpels Syncarpic pistil

Flowers radially Flowers bilaterally

symmetrical symmetrical
Fusion of stamens Staminal tube Fusion of petals Corolla tube
Flowers actinomorphic Flowers zygomorphic
 The Flower The Flower
Adnation:
Adnation: fusion of floral Adnation:
Adnation: fusion of floral
parts from different whorls e.g., Drimys & sandwort parts from different whorls

• Simple adnation
Stamens fused onto inner
surface of fused (connation) No adnation!
petals
Connation (fusion of similar
parts) may or may not occur
• Complex adnation
Sepals, petals, and stamens fuse
to form a hypanthium

The Flower The Flower

Adnation:
Adnation: fusion of floral Adnation:
Adnation: fusion of floral
e.g., cherry & rose parts from different whorls e.g., horse gentian & fuchsia parts from different whorls

Adnation of calyx,
corolla, & stamens
= hypanthium
The Flower The Flower
Floral formula - shorthand notation Floral formula - shorthand notation

4 sepals (CA
(CAlyx)
lyx) 4 sepals (CA
(CAlyx)
lyx)
4 petals (CO
(COrolla)
rolla) 4 petals (CO
(COrolla)
rolla)
8 stamens (A
(Androecium) 8 stamens (A
(Androecium)
4 carpels (G
(Gynoecium) 4 carpels (G
(Gynoecium)
• Carpels fused = 1 pistil

Oenothera biennis Oenothera biennis

Evening primrose Evening primrose
Onagraceae Onagraceae

The Flower The Flower

Floral formula - shorthand notation Floral formula - shorthand notation

4 sepals (CA
(CAlyx)
lyx) 4 sepals (CA
(CAlyx)
lyx)
4 petals (CO
(COrolla)
rolla) 4 petals (CO
(COrolla)
rolla)
8 stamens (A
(Androecium) 8 stamens (A
(Androecium)
4 carpels (G
(Gynoecium) 4 carpels (G
(Gynoecium)
Carpels fused = 1 pistil Carpels fused = 1 pistil
• Ovary inferior Ovary inferior
• Hypanthium (+ hypanthium tube)

Oenothera biennis Oenothera biennis

Evening primrose Evening primrose
Onagraceae Onagraceae
 The “flower” — what is it?
Angiosperms -
• a flower is a specialized shoot that:

1. is determinate (vs. indeterminate)

Floral Structure
2. has a modified stem with compressed internodes

3. possesses modified leaves with various functions, See second half of Chpt 9 for more detail
these determined by gene arrays (e.g., ABC model) (Plant Systematics)
Systematics)
4. often clustered in an inflorescence (larger branch)