Evolution and Diversity of

Land Plants

A. Non-Vascular Plants

Major Reference: Simpson M. Plant Systematics. Elsevier Inc.

 The Land Plants
=Embryophyta (embryophytes)

• The first colonization of plants on land

during the Silurian period, ca. 400 million
years ago.
• Land Plants now dominate the earth.
Figure 29.2

Red algae

ANCESTRAL

Viridiplantae
ALGA Chlorophytes

Streptophyta
Charophytes

Plantae
Embryophytes

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 Chlorobionta - green plants
..---------------....
Embryophytes - land plants* ...-- - -
~ Tracheophytes - vascular plants I
lophytesl r Spennatophytes - see_d plants l
~ rGymnospermsl r Angiosperms 1
=
-= ·--= = -
~
[l:J -
[l:J
[l:J
~
-.
=""'
~ - [l:J
[l:J [l:J
[l:J
,..,I [l:J
·- -0
~
"Green Algae" ,..,I ,..,I "0
0 [l:J 0
[l:J
:.. 't> 0
c:..
,..,I
OJ)
~=
·-= == - c.. -·-= = ·-=
~ 0
[l:J

O"'
:..
:.. c.. .c
....... -~ ,..,I
0
"0
c:..
.......
OJ)
.::-:: ·- 0
=-
0 ,.;
0
=
0 ·-=
c:..
"0
~
[l:J
~ ~ u 0 ~ ~ U5 ~ ~

Flower, carpels,
stamens (+ sev.
other features)

Xylem & phloem vascular tissue

lndependentsporophyte

Cuticle, gametangia, embryo (sporophyte)

Green plant chloroplast

-,- - - - - - - Viricliplantae [Chlorobionta] = Green Plants I
r Chlorophytes 7 Streptophytes - - - - - - - - -

Charophytes
v.,
I
(1)

e Land Plants
c:::
.c:
u = Embryophytes

parenchyma

sporophyte/embryo (alternation of generation )

true starch storage compound

thylakoid stacked in grana Unique green plant chloroplast featur
chlorophyll b (chlorophyll a i ance, tral
 cell wall

chloroplasts

Elodea (Hydrocharitaceae - Alismatales)

 How did chloroplasts evolve?
self.replka ting
chloropl ast
ano stral .
photosy ntheti c
bact rium

eukaryotic ceJJ pbotosyntb tic

ukaryotic ceJJ

….. by endosymbiosis
 Chloroplasts

double {outer membrane

membrane inner membrane

independent overlapping
thylakoids thylakoids
b.

stroma
a.
500nm
-,- - - - - - - Viricliplantae [Chlorobionta] = Green Plants I
r Chlorophytes 7 Streptophytes - - - - - - - - -

Charophytes
v.,
I
(1)

e Land Plants
c:::
.c:
u = Embryophytes

parenchyma

sporophyte/embryo (alternation of generation )

true starch storage compound }

thylakoid stacked in grana Unique green plant chloroplast feat ur
chlorophyll b (chlorophyll a i ance, tral
cellulo e in cell wall (may ha e e oJved earlier & thu not a ynapom rphy for Chlorobionta alone)
 HAPLOID(n) HAPWID(n)
/ Multi lled Sta e / Multicelled Stag
/ (Adult) / ( dull)

@
?~'~ @
(
mitosis

HAPLONTIC HAPLONTIC
@~ Isogamy @~ Oogan1y
Gamet Garn te Spore Egg Sperm
(n) (n) (n) (n) (n)

meiosis @ fertilization meiosis @ fertilization

A ~Zy~,e~
B ~Zygol~

(2n) (2n)
 TYPES OF LIFE CYCLES

DIIPLONTIC HAP LOINTIC HAPLQ,-DIPLQ,NTIC

Ha;ptoid singl1e-cell or Hlap:toi:d multicelllu'lla:r
multicellular oiyantsm organism
Gameres
n Mliitosis
·'!·0
Fe

2n Mitosis
'
Diploid
multicellular
,....._,.~ ' Diploid
multicellular
organism organism
l n animals Most fiun gi
1 Plants
and some algae 1
and :s ome algae

https://www.vcbio.science.ru.nl/en/virtuallessons/lifecycle/
 Charales - oogamy
oogonium
Coleochaete

Nitella Chara
-,- - - - - - - Viricliplantae [Chlorobionta] = Green Plants I
r Chlorophytes 7 Streptophytes - - - - - - - - -

Charophytes
v.,
I
(1)

e Land Plants
c:::
.c:
u = Embryophytes

parenchyma

sporophyte/embryo (alternation of generation )

true starch storage compound }

thylakoid stacked in grana Unique green plant chloroplast feat ur
chlorophyll b (chlorophyll a i ance, tral
cellulo e in cell wall (may ha e e oJved earlier & thu not a ynapom rphy for Chlorobionta alone)
 CELLl C ELL2

pl · ma
~ pla mode mata
membran e middle lamella

cellulosic cell wall

cell wall:
cellulosic
plasmodesmata (in some Green Plants)
-allow cell-to-cell communication
Derived Traits of Plants

Five key traits appear in nearly all land plants but are
absent in the charophytes
a)Alternation of generations
b)Multicellular, dependent embryos
c)Walled spores produced in sporangia
d)Multicellular gametangia
e)Apical meristems

© 2015 Pearson Education Ltd

 mbr ph ta - land plan
Bry

0-
!!l
,> --
~
0
,

=-
C QI i
0
~
.t.

ph t • tin t
nutriti nail

nth ti • nutrili nall ind pen nt

Figure 1. Most recent phylogenetic relationships within the Embryophyta

based both morphological and molecular data sensu Puttick et al. 2018.
Illustration taken from Simpson 2019.

figure 1. Most recent phylogenet ic relat ionships wit hin t he Embryophyta

based bot h morphological and molecular dat a sensu Putticlk et aL 2018 .
Illust rat ion take n from Simpson 2019 .
 Sporophyte Body
HAPLODIPLO TIC
LIFE CYCLE
(" Alternation of Generations")
~
mitosis, growth, & differentiation ------
mitosis, growth, & differentiation
/ ~
mbryo
porangium
/
mitosis, growth, & differentiation
\
mitosis, growth, & differentiation
I

@
Zygote
SPOROPHYTE GENERATION
\
Sporocyte
(2N)

- -
t
fertilization - - - - - - - - - - - __________ _ meiosis

(Spenn non-flagellate in Conifurs , " ~ )) GAMETOPHYTE GENERATION

Gnetales, and Angio perms) ~ ~ (N)

;oresi~
Egg Sperm

lost by reduction and modificatio{ Archegonium Ant.heridium

in the Angiosperms ~ ~ mitosis, growth, & differentiation
and ome Gnetales -.....____ , "'-
mitosis, growth , & differentiation
~Gametophyte Body
/
sporophyte:
- allowed for many more propagules (spores) produced
- sporophyte (diploid) shields against deleterious recessives
m un ·ti nal in
iun

epidermal
 t m

Shoot Tip

parenchyma:
solid mass of tissue; cells:
1) are elongate to isodiametric;
2) have a primary (1˚) cell wall only (rarely a 2˚ wall);
3) living at maturity, capable of continued cell divisions.
Figure 29.3e
Apical meristems
Apical meristem Developing
of shoot leaves

Apical meristem
of root

Root 100 µm Shoot 100 µm

Apical meristems of plant roots and shoots
© 2015 Pearson Education Ltd
 ~por phyt / mbryo alt rnation of enera1

anth ridium neck

par nchyma
uti I
por phyte/ mbry a)t rnati n f r n ration
Table 29.1
Table 29.1 Ten Phyla of Extant Plants

Number
of Known
Common Name Species

Nonvascular Plants {Bryophytes)

Phylum Hepatophyta Liverworts 9,000
Phylum Bryophyta Mosses 15,000
Phylum Anthocerophyta Hornworts 100
--
Vascular Plants
Seedless Vascular Plants
Phylum Lycophyta Lycophytes 1,200
Phylum Monilophyta Monilophytes 12,000
Seed Plants
Gymnosperms
Phylum Ginkgophyta Ginkgo 1
Phylum Cycadophyta Cycads 130
Phylum Gnetophyta Gnetophytes 75
Phylum Coniferophyta Conifers 600
Angiosperms
Phy Ium Anthophyta Flowering plants 250,000
© 2015 Pearson Education Ltd
Figure 29.5

ANCESTRAL
Liverworts

Land plants
(bryophytes)
plants
Nonvascular
GREEN
ALGA Origin of land plants
1 Mosses

Hornworts

Lycophytes (club mosses,

Vascular plants
plants
vascular
Seedless
spike mosses, quillworts)
Origin of vascular plants
2 Monilophytes (ferns,
horsetails, whisk ferns)
Gymnosperms

plants
Seed
Origin of extant
3 seed plants
Angiosperms
(
500 450 400 350 300 50 0
Millions of years ago (mya)

© 2015 Pearson Education Ltd

Hornworts

Mosses

Liverworts

BRYOPHYTES

E. Mandia 12/6/2013; DLSU-STC

 Bryophyte diversity
Lumpers’ notion:
Division Bryophyta
3 classes (Musci, Hepaticae, Anthocerotae
[ICN: Bryopsida, Hepatopsida,
Anthoceropsida])
Splitters’ notion:
Division I : Bryophyta (Mosses)
Division II: Hepatophyta (Liverworts)
Division III: Anthocerophyta (Hornworts)
r:=:=======~ J.!!!!~O
;!!P!!h~y~ta~ - land plants - - - - - - -
Pan-Tracheophyta/
Polysporangiophyte
I I
Tracheophyte
t t -vascular plant,

gametophytic
pseudo-elaters
l aves in sporangium

columella in
sporangium sporophyte branched
leptoids with multiple porangia
intercalary growth
of sporophyte
hydroids
oil bodie
porophyte photo ynthetic, nutritionally independent

aerial sporoph yte axis

tomates

ar hegonium t = extinct
antheridi um
parenchyma
“Bryophytes” paraphyletic
porophyte/embryo (alternation of generation )
 mbr oph ta - land plan

,
C
L,,

=
0
::s

ph 1· . tin t
nutriti nall

ia
p ud
m

mat· • un ti nal in po n ium han

[alt m li

-fun ti n, I in poran ium d hi Monophyletic?

B Figure 1. Most recent phylogenetic relationships within the Embryophyta
based both morphological and molecular data sensu Puttick et al. 2018.
n.
Illustration taken from Simpson 2019.
THE HORNWORT Anthoceros
Mature
sporangium
• splits open
• . •
• • .
• Spores
. .• u

Sporophyte

E. Mandia 12/6/2013; DLSU-STC

Gametophyte with
embedded archegonia
and antheridia
A) Habit of Anthoceros (b) After fertil ization , the sporophytes project up out of the
gametophyte thal lus, forming "horns ."
@ 2007 Thomson Higher Education
 The Hornworts
Thalloid body
• Single large chloroplast per cell
– Gametophyte
• Discoid, photosynthetic
– Sporophyte
• Photosynthetic, with stomata
• Long and narrow, splits lengthwise
• Indeterminate growth due to intercalary meristem
• With columella and pseudo-elaters in the sporangium
– primarily tropical in distribution
Hornwort
sporophyte ls
 chi , pla! u

stomata
:t }m,,

Stomata in hornwort
 Liverworts
Division Hepatophyta

8000 species worldwide

2 orders:
1. Marchantiales
(“Thalloid”)
E. Mandia 12/6/2013; DLSU-STC

2. Jungermanniales
(“Leafy”)
 Leafy (Bazannia)
-3 rows of leaves,
lower usu. reduced
Asexual reproduction in the Liverworts
is through formation of
gemmae cups
gemmae
 Liverwort forms Gametangia

Antheridiophore

Gametophyte thallus

(a) Antheridiophores produce sperm-bearing antheridia.

@ 2007 Thomson Higher Education
 Sporophyte

Archegoniophore

(b) Archegoniophores produce egg-bearing archegonia.

After fertilization of the egg within an archegonium, the
sporophyte generation develops. It hangs upside down
under the fingerlike projections of the archegoniophore
and resembles a miniature coconut hanging on a palm tree.
@ 2007 Thomson Higher- Education
 egg cell

neck

archegonium

Marchantia (liverwort)
Sporophyte of Marchantia
 Antheridiophore
Archegoniophore
Male
Germination of spores
thallus 1
and development of
young gametophyte Female
5 thallus Antheridia with
Gemmae sperm cells
cup
Spores Male and female Sperm
released gametophyte plants cell

HAPLOID (n) Archegonia

GAMETOPHYTE with eggs 2
GENERATION
Meiosis Fertilization
DIPLOID (2n)
4 SPOROPHYTE
GENERATION
Foot
Seta
Zygote
Tissue derived
from archegonium
Capsule Embryo
Spore
mother cells that
undergo meiosis 3
Sporophyte

© 2007 Thomson Higher Education

Fig. 22-9, p. 439
 Mosses
(Division Bryophyta)

sporophyte

gametophyte
 Bryophyte diversity
Division I : Bryophyta (Mosses) -12,000 species
• Class Sphagnopsida (peat mosses: 200 Sphagnum spp.)
•about 1% of earth terrestrial surface covered by Sphagnum

• Class Andreaeopsida (lantern mosses)

• Class: Polytrichopsida
• Class Bryopsida (95% of moss species)
 Sphagnum - Peat moss:
Plants make ground water acidic, form extensive bogs

Sphagnum Peat Moss

 Peat = partially decomposed
Sphagnum

- Used in
potting media

- Used as a
fuel
 Peat = partially decomposed Sphagnum

- “Tolland” man:
- preserved
remains in bog,
405-100 B.C.

"Tolland Man," a bog mummy dating from 405-100 B.C.

The acidic, oxygen-poor conditions produced by Sphagnum can
preserve human or other animal bodies for thousands of years.
R1mrly !Moore,. Den nis Oledi,. and Dslrell Vodopich, Bo!BD!f' II uisl Resource Ubraiy 1998 'll'he McGlillW-HUI Oampemi~, lne. ll ri.9J}ts r@!l@f'i'Bd

Operculum-
Structure of Mosses

Sporophyte :. ~porogenous
(2n) tissue
(2n)

Gamet,ophyte
(n)

(a)
 operculum pecistome teeth

spores
peristome
tooth

sporophyte
/ (2n)

gametophyte
(n)
spore protonema gametophyte (n)
(n)
meiosis

fertilization

sporophyte (2n) archegonium antheridium

Figure 29.6

“Bud”
Sperm
Protonemata
(n) Antheridia I
_I_I
Key Male
gametophyte (n)
•• Haploid (n)
Diploid (2n) “Bud”

Egg

Spores Gametophore
Spore Archegonia
dispersal
. .... . ...
Female Rhizoid
Peristome gametophyte
Sporangium (n)

Seta FERTILIZATION
MEIOSIS Capsule (within archegonium)
(sporangium) Zygote
Mature (2n)
Foot Embryo
sporophytes

Archegonium
2 mm

Young
Capsule with Female sporophyte
peristome (LM) gametophyte (2n)
© 2015 Pearson Education Ltd
The Ecological and Economic Importance of Mosses
1. Mosses are capable of inhabiting diverse and sometimes extreme
environments, but are especially common in moist forests and
wetlands
a) important role as pioneer organisms
2. Some mosses might help retain nitrogen in the soil
3. Sphagnum is an important global reservoir of organic carbon
a)Peat can be used as a source of fuel
b)Low temperature, pH, and oxygen level of peatlands inhibits decay
of moss and other organisms
c)Overharvesting of Sphagnum and/or a drop in water level in
peatlands could release stored CO2 to the atmosphere

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Thank you for your attention