Chapter 21

Lecture Outline

The Seedless
Vascular Plants:
Ferns and Their
Relatives

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• Introduction Outline
• Phylum Psilotophyta – The Whisk Ferns
• Phylum Lycophyta – The Ground Pines, Spike
Mosses and Quillworts
• Phylum Equisetophyta – The Horsetails and
Scouring Rushes
• Phylum Polypodiophyta – The Ferns
• Fossils
 Introduction
• During early stages of vascular plant evolution:
– Internal conducting tissue developed.
– True leaves appeared.
– Roots that function in absorption and anchorage
developed.
– Gametophytes became progressively smaller.
• Four phyla of seedless vascular plants:
Psilotophyta, Lycophyta, Equisetophyta,
Polypodiophyta
• Psilotophyta Introduction
– Sporophytes have neither
true leaves, nor roots.
– Stems and rhizomes fork
evenly.
Psilotum
• Lycophyta
– Plants covered with
microphylls.
• Microphylls - Leaves with single
vein whose trace is not
associated with a leaf gap
Lycopodium
• Equisetophyta Introduction
– Sporophytes have ribbed
stems containing silica.
– Have whorled, scalelike
microphylls that lack
chlorophyll Equisetum
• Polypodiophyta
– Sporophytes have megaphylls
that are often large and much
divided.
• Megaphylls - Leaves with more
than one vein and a leaf trace
associated with leaf gap
A fern
• Lycophyta
– Plants covered with microphylls.
• Microphylls - Leaves with single
vein whose trace is not
associated with a leaf gap

• Polypodiophyta Lycopodium
– Sporophytes have
megaphylls that are often
large and much divided.
• Megaphylls - Leaves with
more than one vein and a
leaf trace associated with
leaf gap

A fern
 Phylum Psilotophyta – The Whisk
 Resemble small, green whisk brooms
Ferns
• Structure and form:
– Sporophytes:
• Dichotomously forking stems
– Above ground stems arise from
rhizomes beneath surface of
ground.
• Have neither leaves nor roots
• Enations along stems.
– Enations - Tiny, green,
superficially leaflike, veinless,
photosynthetic flaps of tissue
• Roots, aided by mycorrhizal fungi,
scattered along rhizomes.
 Phylum Psilotophyta – The Whisk
• Reproduction: Ferns
– Sporangia fused in threes and produced at tips of
short branches.
– Gametophytes develop from spores beneath ground.
• Branch dichotomously
• No chlorophyll
• Rhizoids aided by mycorrhizal fungi.
• Archegonia and antheridia scattered on surface.
– Zygote develops foot and rhizome.
– Rhizome separates from foot.
 Phylum Psilotophyta – The Whisk
• Reproduction: Ferns
 Phylum Psilotophyta – The Whisk
Ferns
• Fossil whisk fern look-alikes:
– Silurian, 400 million years ago
• Cooksonia and Rhynia
– Naked stems and terminal sporangia
– Devonian, 400-350 million years ago
• Zosterophyllum
– Naked stems and rounded sporangia along stem
– Thought to be ancestral to club mosses
 Phylum Lycophyta – The Ground Pines,
Spike Mosses, and Quillworts
• Collectively called club mosses
– Two living major genera
• Lycopodium
• Selaginella
– Two living minor genera
– Several genera that became extinct about 270 million
years ago
• Sporophytes have microphylls.
• Have true roots and stems
•
Phylum
Lycopodium -
Lycophyta
Ground pines
– Often grow on forest
floors
– Stems are simple or
branched.
• Develop from
branching rhizomes
– Leaves usually less
than 1 cm long.
– Roots develop along
rhizomes.
 Phylum Lycophyta
 Lycopodium reproduction:
– Sporangia in axils of
sporophylls.
• Sporophyll - Sporangium-
bearing leaves
• In some species,
sporophylls have no
chlorophyll, are smaller
than other leaves and
clustered into strobili
(singular: strobus).

• In sporangia, sporocytes undergo meiosis,

producing spores.
 Phylum Lycophyta
• Lycopodium reproduction:
– Spores grow into independent
gametophytes.
• In some species, gametophytes
resemble tiny carrots, develop in
the ground and are associated
with mycorrhizal fungi.
• In others, gametophytes develop
on surface and are green.
• Archegonia and antheridia
produced on gametophytes.
• Sperm are flagellated and water is
essential for fertilization.
 Phylum Lycophyta
• Lycopodium
reproduction:
 Phylum Lycophyta
• Selaginella - Spike mosses
– Especially abundant in tropics
– Branch more freely than ground pines
– Leaves have a ligule on upper surface.
•
Phylum Lycophyta
Selaginella reproduction:
– Produce two different kinds of gametophytes =
heterospory.
• Microsporophylls bear microsporangia containing
microsporocytes, producing tiny microspores.
– Microspore becomes male gametophyte, consisting of an
antheridium within microspore wall.
• Megasporophylls bear megasporangia containing
megasporocytes, producing 4 large megaspores.
– Megaspore develops into female gametophyte consisting of many
cells inside megaspore.
– Several archegonia produced where spore wall ruptures.
 Phylum Lycophyta
• Selaginella reproduction:
 Phylum Lycophyta
• Isoetes - Quillworts
– Most found in areas partially submerged in water for
part of year.
– Microphylls are arranged in a tight spiral on a stubby
stem.

• Ligules occur
towards leaf bases.
• Corms have
vascular cambium.
• Plants generally
less than 10 cm tall.
 Phylum Lycophyta
• Isoetes
reproduction:
– Similar to spike
mosses, except no
strobili
– Sporangia at bases
of leaves.
 Phylum Lycophyta
• Ancient relatives of
club mosses and
quillworts:
– Dominant members
of forests and
swamps of
Carboniferous, 325
million years ago
• Large, tree-like, up to
30 meters tall -
Lepidodendron

Surface of Lepidodendron,
showing microphyll bases
 Phylum Equisetophyta – The Horsetails
and Scouring Rushes
• Equisetum
• Branched and unbranched
forms, usually less than 1.3
meters tall
• Stems jointed and ribbed.
– If branched, then branches in
whorls.
– Scalelike leaves in whorls at
nodes.
– Stomata in grooves between
ribs.
 Phylum Equisetophyta
• Stem anatomy:
– Hollow central cavity from break down of pith
– Two cylinders of smaller canals outside pith.
– Carinal canals
conduct water
with xylem
and phloem to
outside.
– Vallecular
canals outside
carinal canals
contain air.

 Silica deposits on walls of stem epidermal cells.

 Phylum Equisetophyta
• Equisetum reproduction:
– Asexual by fragmentation of
rhizomes
– Sexual reproduction:
• Strobili at tips of stems with
sporangia connected to
sporangiophores.
• Spores green with 4 ribbon-
like elaters attached.
– Aid in spore dispersal
• Gametophytes lobed, green,
cushionlike, up to 8 mm in
diameter.
Spores with
elaters
 Phylum Equisetophyta
• Equisetum reproduction:
 Phylum Equisetophyta
• Ancient relatives of horsetails:
– Flourished in Carboniferous, 300
million years ago.
• Human and ecological relevance:
– Many giant horsetails used for
food by humans and other
animals.
– Scouring rush stems used for
scouring and sharpening.

Reconstruction of fossil giant

horsetail, Calamites
 Phylum Polypodiophyta – The Ferns
• Structure and form:
– Vary in size from tiny floating forms less than 1 cm to
giant tropical tree ferns up to 25 m tall
• Fern leaves are megaphylls - Referred to as fronds.
– Typically divided into smaller segments
• Require external water for reproduction
 Phylum Polypodiophyta – The Ferns

• Reproduction:
– Sporophyte is conspicuous
phase.
• Fronds, rhizomes, roots
Crozier
• Fronds first appear coiled in
crozier (fiddlehead), and
then unroll and expand.
– Fronds often divided into
segments called pinnae
(singular: pinna).
 Phylum Polypodiophyta – The Ferns
• Reproduction:
– Sporangia stalked.
• May be scattered on lower
leaf surface, confined to
margins, or found in discrete
clusters called sori (singular:
Sorus covered by
sorus). indusium
– Sori may be protected by
indusia (singular: indusium).
• With row of heavy-walled,
brownish cells = annulus
– Annulus catapults spores out
of sporangium.
 Phylum Polypodiophyta – The Ferns
• Reproduction:
– Meiosis forms spores in sporangia.
– Spores released and grow into gametophytes called prothalli
(singular: prothallus).

• Prothalli are one cell thick,

and have archegonia and
antheridia.
• Zygote develops into
young sporophyte.
• Gametophyte, or portion of
it, dies and leaves
sporophyte growing
independently.
 Phylum Polypodiophyta – The Ferns
• Reproduction:
 Phylum Polypodiophyta – The Ferns
• Fossil relatives of ferns:
• Devonian, 375 million
years ago - Possible
ancestors of ferns
– Resemble ferns in
growth habit, but look
more like whisk ferns

Possible ancestors: Aglaophyton and Psilophyton

 Phylum Polypodiophyta – The Ferns
• Fossil relatives of ferns
– Carboniferous, 320-250
million years ago - Tree
ferns abundant
• Seeds found on some of
fossil tree ferns.
•
Phylum Polypodiophyta
Human and ecological relevance:
– House plants
• Function well as air filters
– Outdoor ornamentals
– Cooked rhizomes as food
– Folk medicine
– Fronds used in thatching for houses.
– Basketry and weaving
 Fossils
• A fossil - Any recognizable prehistoric organic
object preserved from past geological ages.
– Conditions of formation almost always include quick
burial and an accumulation of sediments.
• Hard parts more likely preserved than soft parts.
 Fossils
• Molds, casts, compressions, and imprints:
– After being buried in sediment and hardened into rock,
organic material slowly washed away.
– If air space remains - Mold
– If silica fills space - Cast
• Compression - Objects buried
by layers of sediment and
overlying sheer weight
compresses them to thin film of
organic material and an outline.
– Image of an impression = imprint
– Coal is a specific type of
compression.
Compression fossil
 Fossils
• Petrifactions - Uncompressed rock-like material in which
original cell structure has been preserved
– Chemicals in solution infiltrate cells and cell walls, where they
crystallize and harden, preserving original material.

 Coprolites - Dung of
prehistoric animals
and humans
 Unaltered fossils -
Organisms fell into
oil or water that
lacked oxygen and
did not permit decay. Petrified wood
• Introduction Review
• Phylum Psilotophyta – The Whisk Ferns
• Phylum Lycophyta – The Ground Pines, Spike
Mosses and Quillworts
• Phylum Equisetophyta – The Horsetails and
Scouring Rushes
• Phylum Polypodiophyta – The Ferns
• Fossils