Chapter 4: Evolution and

Diversity of Vascular Plants

Michael G. Simpson
Vascular Plants = Tracheophyta
(Tracheophytes)
Alternation of Generations: Haploid (n) & Diploid (2n) adult phases

Bryophytes: Gametophyte
dominant, long-lived
Bryophytes (Liverworts, Hornworts, Mosses):
Gametophytes are dominant, long-lived
Alternation of Generations: Haploid (n) & Diploid (2n) adult phases
Vascular Plants:
Sporophyte dominant, long-lived
Vascular Plants: Sporophytes are dominant
Sporophytic axes
branched with
multiple sporangia

Sporophyte of a moss
Rhynia
Two early branching patterns in
vascular plants:
 lignin - hard substance secreted
within secondary cell wall

Major adaptive:
added structural support enabled
vascular plants to grow much larger.
 secondary cell wall

- forms between 1 cell wall and plasma

membrane in some plant cells

- adds structural support

- found in tracheids, vessels, fibers

(all dead cells)
 Primary cell wall: cellulosic
lignified
secondary cell
wall
Sclerenchyma
1) Secondary cell wall (+ primary)
2) Dead at maturity (usually)

Fibers
Elongate, sharply tapering
Sclerenchyma
1) Secondary cell wall (+ primary)
2) Dead at maturity (usually)

Sclereids
Isodiametric to irregular
 Tracheary elements
1) cells that function in water/mineral conduction
2) cells dead at maturity, with lignified, 2 cell walls
3) cells arranged end-to-end, forming long tubes

Xylem = tissue composed of:

1) tracheary elements
2) parenchyma
3) fibers
(These have common origin/location/function.)
 Tracheary Elements
Tracheids imperforate (only pits at end Vessels perforation plates = holes in end walls.
walls). Found in most monilophytes (except Evolved independently from preexisting
Equisetum and some lepto. ferns) and most tracheids in Equisetum, a few leptosporangiate
gymnosperms (except Gnetales) ferns, all Gnetales, and almost all angiosperms
 Sieve Elements
-cells with sieve areas/plates, having pores lined by callose
callose (polysaccharide composed of -1,3-glucose units)
Sieve cells only sieve areas Sieve tube members - Sieve plates
All vasc. pls. except angiosperms Apomorphy of Angiosperms
 Phloem
= tissue composed of:
1) sieve elements
2) parenchyma
3) fibers

These have common origin/function.

 Endodermis single layer of cells surrounding
vasculature of roots and some underground stems.

Casparian Strip: forces fluids from outside through

plasma membrane = selective absorption
 Roots
Function in
anchoring and
absorption.
 Root

apical meristem
- region of actively
dividing cells
 Roots
Five diagnostic features:
1) Root cap functions to protect apical meristem
and lubricate root during growth.
2) Root hairs function to greatly increase
water/mineral absorption (exception Psilotopsida).
3) Central vascular cylinder site of conduction of
water/minerals and sugars.
4) Endodermis Casparian strip functions in
selective absorption.
5) Endogenous lateral roots develop from
pericycle or endodermis, penetrate to outside.
 Mycorrhizae
Symbiotic association with various species of fungi.
Found in many, if not most, vascular plants

Fungus aids plant by

1) increasing surface area for water and mineral
absorption
2) increasing efficiency of mineral absorption.

Fungus benefits in obtaining photosynthates (sugars and

other nutrients) from the plant.
Tracheophyta Classification
 Lycopodiophyta lycophytes

Apomorphies:
1) Roots dichopodial
-apical meristem
may branch into two.

2) Root protoxylem endarch (to inside)

Protoxylem first xylem formed
Metaxylem later xylem
3) Stem protoxylem exarch (to outside)
 4) Sporangia dorsiventral
(flattened upper and lower surfaces)
with transverse dehiscence.
Shoot = stem + leaves
Lycophyll simple leaf with 1 vein, no leaf gaps.
Tracheophyta Classification
LYCOPODIOPSIDA
Lycopodiaceae
homosporous = 1 type of
spore
5 genera: Huperzia,
Lycopodiella, Lycopodium,
Pseudolycopodiella,
Phylloglossum)

Lycopodium digitatum
 Lycopodium spores:
Used in original camera flash powders,
even some fireworks
 ISOETOPSIDA
Selaginellaceae + Isoetaceae
1) Leaves ligulate
2) Heterosporous 2 types of spores

Ligule tiny appendages on upper side of leaf (near base)

Heterospory two types of spores:
megaspores (from megasporangia)
microspores (from microsporangia)
Selaginellaceae
Spike-moss
family

1 genus:
Selaginella
(700 spp.)

Selaginella
spike-moss
Selaginella bigelovii
Selaginella cinerascens
 Some
Selaginella
have
dimorphic
leaves 2 rows large
leaves

Selaginella apoda
 Some
Selaginella
have
dimorphic
leaves
2 rows small
leaves

Selaginella apoda
 Isoetaceae
Quillwort family

1 genus:
Isoetes (ca. 200 spp.)

Isoetes quillwort/Merlins grass

Lycopods native to San Diego
area:
Isoetes Merlins Grass
Isoetes orcuttii

Isoetes howellii
 Extinct Lycopods were
tall trees:
make up some of coal
deposits

Lepidodendron
Euphyllophyta

Apomorphies:
1) roots monopodial
2) root protoxylem exarch
3) sporangia terminal on
lateral branches,
longitudinally dehiscent

Archeopteris
Euphyllophyta

Apomorphies:
1) roots monopodial
2) root protoxylem exarch
3) sporangia terminal on
lateral branches,
longitudinally dehiscent
4) shoot with euphylls

shoot = stem + leaves

Euphyllophyta

Apomorphies:
1) roots monopodial
2) root protoxylem exarch
3) sporangia terminal on
lateral branches,
Leaves of euphyllophytes:
longitudinally dehiscent
a) evolved by planation & webbing of
4) shoot with euphylls stems
b) have multiple veins
c) have a leaf gap
Euphyllophyta

Apomorphies:
1) roots monopodial
2) root protoxylem exarch
3) sporangia terminal on
lateral branches,
longitudinally dehiscent
4) shoot with euphylls
5) 30 kb chloroplast
DNA inversion
Tracheophyta Classification
Monilophyta
-monilophytes

Apomorphies:
1) siphonostele
2) stem protoxylem
mesarch
 Equisetopsida
(Equisetophytes)

Equisetaceae
Equisetum Horsetails
/ Scouring Rushes
Equisetopsida

Apomorphies:
1) ridged stem
with canals
2) leaves reduced,
whorled
Equisetopsida

Apomorphies:
1) stem ribbed
with canals
2) leaves reduced,
whorled
3) sporangiophore
4) spores with elaters,
chlorophyllous
 cone
(strobilus)

whorled
microphylls

Equisetum hyemale common scouring-rush Equisetum laevigatum smooth scouring-rush

Subgenus Hippochaete scouring-rushes

Equisetum arvense Common Horsetail

Subgenus Equisetum horsetails

 Calamites

-fossil (extinct)
equisetophyte,
was tree-sized, makes
up coal deposits today
Psilotopsida
Apomorphies:
1) Roots unbranched,
root hairs absent.
2) Gametophyte
subterranean,
mycorrhizal.
Psilotopsida
Two members:

1) Ophioglossales
ophioglossoid ferns

2) Psilotales
whisk ferns
Ophioglossales:
One family: Ophioglossaceae
4 genera, incl. Botrychium, Ophioglossum
Ophioglossaceae
Ophioglossum californicum Calif. Adders Tongue
Psilotales

Apomorphies:
1) Roots lost.
2) Leaves reduced
(microphylls or
enations).
3) Synangium with bifid
appendage.
Psilotales

One family:
Psilotaceae
2 genera:
Psilotum
Tmesipteris
dichotomous
branching

Psilotum nudum
Marattiopsida
Marattioid Ferns

Apomorphy:
1) Polycyclic
siphonostele
Marattiaceae

eusporangia
Polypodiopsida
Leptosporangiate Ferns

Apomorphy:
Leptosporangium
Polypodiopsida
Leptosporangium:
- one cell thick
- spores ejected
Stem:
rhizome
trunk
vine
fern leaves can have specialized terminology:
leaf = frond
petiole = stipe
major divisions = pinna (1, 2, etc.)
ultimate divisions = pinnules
 fern leaf venation:
open (simple or forked), reticulate

fern scales:
e.g., clathrate or non-clathrate
sorus (pl. sori) aggregation of sporangia
indusium - flap

shape /
morphology
 leptosporangium / annulus
-varies with different groups/families
sperm

egg
 Fern leaf development: circinate vernation

fiddle head - crozier

 Salviniales
Apomorphies:
1) aquatic adaptation
2) heterospory (independently evolved)
3) sporocarps (dormant, seed-like body,
house specialized sporangia)

Two famlies:
Marsileaceae clover fern family
Salviniaceae floating fern family
Marsilea Salvinia Azolla
Marsileaceae

Marsilea

Pilularia
Salviniaceae

Azolla

Salvinia
Azolla Mosquito Fern
- symbiotic relationship with blue green bacteria; seeded in rice
paddies for nitrogen fixation
- inhibits mosquito population by covering surface
- fodder for animals
 Cyatheales:

Cyatheaceae

largest family of
tree ferns
(trunk arborescent)
 Polypodiales:
A few
families:
Aspleniaceae

-linear sori /
indusia

Cyathium
a tree fern
Polypodiaceae

- sori exindusiate
(naked)
Pteridaceae
-exindusiate
-false indusium or
intramarginal (in
lines along veins)

Cyathium
a tree fern