Chapter 5


Evolution of Woody and Seed Plants

Michael G. Simpson
Evolution of wood 

= secondary xylem 

derived from vascular cambium
enabled plants to grow very tall,
as large trees.

Illustrated: Sequoiadendron giganteum, giant

sequoia, largest non-clonal organism on
earth; most tissue dead secondary xylem
(wood).
Divisions of vascular cambium mostly tangential (some radial)
2˚ xylem –> inside, 2˚ phloem –> outside: bifacial growth.
 Secondary growth from lateral meristems: 

vascular cambium & cork cambium.

Vascular cambium: 2˚ xylem –> inside; 2˚ phloem –> outside: 


= bifacial growth.
Tissue to outside (cortex, older 2˚ phloem) gradually crushed.
In most temperate regions (growth season / winter) and some tropical
regions (dry/wet season), seasonal annual rings form.
Annual rings can be used to date wood.
Dendrochronology – study of wood in relation to time and environment/
climate.
cork cambium
- like vascular cambium – bifacial
cork –> outside, phelloderm –> inside
 Seed

= embryo enclosed by nutritive tissue,
surrounded by seed coat

Ovule

= immature seed (prior to fertilization/
maturation)
Seed Evolution
Heterospory
 telomes & lagenostome – found in micropyle – hole in integument, 

fossil (extinct) taxa, functioned in pollen where pollen enters (gymnosperms)
capture

pollination droplet – secreted from

micropyle; pollen grains stick to, are
pulled inside as droplet evaporates.

Ginko biloba
 Pollen grains = immature endosporic male gametophytes

Pollen grains develop from microspores by mitotic divisions/differentiation.

[When haploid nucleus of microspore starts to divide mitotically, it becomes a pollen grain.]
 Pollen tube – exosporic, tube-like extension from
pollen grain.

Pollen tube – haustorial (parasitic, feeding off tissues) in Gymnosperms

-In cycads & Ginkgo sperm delivered to fertilization chamber, where sperm
swims to archgonium = zooidogamy.
-In conifers (incl. Gnetales) tube grows directly to archegonium =
siphonogamy.
Ovule development
 Adaptive significance of seeds

1) Protection of embryo (seed coat)

2) Nutrition of embryo (nutritive tissue=female
gametophyte in Gymnosperms)
3) Dispersal unit (e.g., fleshy, carried by animals; winged,
transported by wind)
4) Dormancy mechanisms (seed coat involved in
preventing germination except when conditions right)
eustele – single ring of vascular bundles
 Two fossil (extinct) lignophytes:

Archeopteris – large tree, wood like Medullosa – a “tree fern,” bore seeds, but with
fern-like foliage.
a conifer, leaves like a fern; some
heterosporous.
Gymnospermae – (“naked seed”) seeds not enclosed by carpel; 

sister to Angiospermae.
Cycadophyta - Cycads
 Cycadophyta – Cycads



-trunks short (rarely elongate, tree-like)











-leaves pinnate (rarely bipinnate), 

coriaceous, with circinate 

vernation (like ferns)











-dioecious (separate male & female 

individuals)







-sperm motile
 Cone = determinate shoot systems, consisting of a single axis with
sporophylls = modified leaves with attached sporangia.

Pollen cones (male) – bear

microsporophylls, with
microsporangia.
pollen cone

seed cone

Ovulate / seed cones

(female) – bear
megasporophylls, 

with ovules/seeds.
 Cycadophyta – Cycads



Cycadaceae: 

female plants 

without cones; 

seeds born on 

megasporophylls 

from stem axis.



One genus: Cycas



Cycas revoluta

sago-palm 

-starch derived from

pith –> flour, bread.

[C. media

-edible seeds]
 Cycadophyta – Cycads



Zamiaceae: 

female plants 

with seed 

cones; 

leaves pinnate 

or bipinnate
 Cycadophyta – Cycads



Zamiaceae: 

female plants 

with seed 

cones; 

leaves pinnate 

or bipinnate
 Ginkgophyta – Maidenhair Trees

One family:
Ginkgoaceae
Maidenhair Tree family

One species
(monotypic):
Ginkgo biloba
 Ginkgophyta - Ginkgo biloba


 - venation dichotomous

- tree with short shoots

- dioecious
- leaves obtriangular
 Ginkgo biloba
Pollen cones “catkin-like”: axis bearing stalk-like microsporophylls, each with
two microsporangia.
 Ginkgo biloba
Ovulate reproductive structures: stalk bearing two, erect ovules, each with basal
collar. Seeds fetid (butyric acid).
 Coniferae – conifers

Apomorphy:
pollen tube
– siphonogamous
-leaves simple

Three groups:
Pinopsida
Cupressopsida
Gnetales
Pinaceae: Pinus
Mystery of the conifer cone: compound
 Evolution of the conifer cone:
Compound structure (axis bearing two components):
bract – homologous with leaf
ovuliferous scale – homologous with shoot system
 Coniferae – conifers

Pinopsida:

One family:
Pinaceae
Pinaceae

-pollen cones with

2 microsporangia
/ microsporophyll

-pollen grains of
some taxa saccate
(with 2 bladders)
Pinaceae

-seeds usu. 2 per

ovuliferous scale,
inverted, winged.
Pinaceae
-leaves linear to acicular
(needle-shaped)

- in some taxa, 

short shoots 

(e.g., Cedrus)
or 

determinate fascicles
(Pinus).
Pinaceae
Include:
Abies – fir
Cedrus – cedar
Larix – larch
Pinus – pine
Pseudotsuga –Douglas-fir
Tsuga - hemlock
Pinaceae
Includes oldest non-clonal
organism on earth:
Pinus longaeva,
bristlecone pine,
5,063 years
old!
 Coniferae – conifers

Cupressopsida

5-6 families
We will cover 4:
Araucariaceae
Podocarpaceae
Cupressaceae
Taxaceae
Araucariaceae

-leaves broad to acicular

-microsporangia
numerous (5-20) per
microsporophyll

-ovule 1 per scale

Includes:
Araucaria heterophylla
Norfolk Island-Pine
Araucaria bidwillii
bunya-bunya
Agathis australis
kauri
Cupressaceae

-leaves linear, acicular, or

subulate, spiral, opposite,
or whorled
-branches flattened in some,
resembling pinnate leaves
in some
-seed cones with numerous
seeds per scale (2-20)
-pollen not saccate

Includes:
Cupressus – cedar
Juniperus - juniper
Sequoia sempervirens –
redwood
Sequoiadendron giganteum –
giant sequoia
Cupressaceae

Includes most massive, non-

clonal organism on earth:
Sequoiadendron giganteum –
giant sequoia

more massive than a blue

whale (largest animal) or any
dinosaur that ever lived




Cupressaceae





and tallest living tree in the
world:
Sequoia sempervirens –
redwood,

Tallest measured (to date) 


at 379.1 feet (115.5 m).
Podocarpaceae

-leaves linear, elliptic, or

subulate
-seed cones usu. fleshy, often
reduced, in some subtended
by fleshy receptacle, seed
may be enveloped by fleshy
epimatium derived from
scale

Includes:
Podocarpus, e.g., P. gracilior
Phyllocladus spp.
Taxaceae
Yew family

-seed cones usu. reduced

to 1 ovule
-seed surrounded by
fleshy aril

Includes:
Taxus – yew
taxol derived from T.
brevifolia, used to treat
ovarian cancer
 Coniferae – conifers

Gnetales:

3 families:
Ephedraceae
Gnetaceae
Welwitschiaceae

All dioecious
Gnetales:

Apomorphies:
1)Pollen striate
2)Vessels porate
 Gnetales

Gnetaceae

One genus: Gnetum (ca. 30 spp.)

Gnetum – tropical vines [trees, shrubs], 


leaves simple, opposite
Gnetales

Ephedraceae



One genus:

Ephedra (35-45 spp.)

Morman Tea



E. sinica:

ma-huang – Chinese

herbal medicine



ephedrine – alkaloid

weight loss, stimulant

now banned.
 Ephedra Morman-Tea
Shrubs
Deserts of S.W. North America, 

W. South America, N. Africa, and Eurasia
Ephedra



- pollen cones
with stalk-like
microsporan-
giophore,
bearing
synangia



- seeds of seed
cones with
fleshy, connate
bracteoles and
micropylar tube
 Gnetales

Welwitschiaceae



One genus/species:

Welwitschia mirabilis
-native to deserts 

of Namibia