Sources: Freeman (2002), Purves et al (2001)

Evolution of Land Plants

Prior to the origin and diversitication of green plants in the mid-Silurian
(~450 mya), multicellular life was virtually entirely adapted to, and
confined to, aquatic lifestyles
Continental land Masses virtually unoccupied by multicellular organism –
multicellular-based ecosystems constituted tremendous potential for
adaptive radiation
Terrestrial life in a gaseous medium required evolutionary solutions to
structural, physiological and ecological challenges; Many of these
innovations can be regarded as exaptations of pre-existing traits of green
algae from which green plants diverged

Artist’s rendering of Carboniferous forest in a tropical river delta. Most of

the plants depicted here were nonseed tracheophytes 10 to 20 meters tall.
In the distance, early seed plants up to 40 meters tall towered over the
forest.
 Sources: Freeman (2002), Purves et al (2001)

Kingdom Plantae is monophyletic assemblage,

descended from Green algae
Chlorophyll a and b are homogous in Chlorophyta,
Charaphyta and Plantae
Currently includes over 250,000 described species
classified in 12 monophyletic phyla
Most are terrestrial, some are secondarily aquatic
Diversification in plants involved successive
adaptive radiations following evolution of key
innovations that increased efficiency in a gaseous
Ie, “true”
vasular
(air) and solid (Earth) environment
tissue Major problems were posed by gravity and by
water loss/availability
-maintain body structure in air (and with
increasing body size) – resist gravity
-obtain, transport and retain water
-fertilize eggs and produce and protect
embryos
Major evolutionary innovations included
-dimorphic body
-waxy cuticle and stomata
- true vascular tissue;
-life history dominated by sporophyte
generation
-jacketed sex organs; antheridia and -
archegonia
-seeds; embryo with nutritive tissue in
Green algae of the lineage that includes Chara sp., are protective covering
most likely ancestors of plants. Copyright BPS. -flowers; vehicles for pollination strategies
 Sources: Freeman (2002), Purves et al (2001)
Sources: Freeman (2002), Purves et al (2001)
20
First extensive
grassland

140
First Flowering
Plant

First nectar-drinking
insects

First vessels
280
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

1st seeds
360
1st vascular plant leaves
1st wood
1st roots
1st vascular tissue
410 1st stomata
1st megafossils (small club
mosses and other extinct
groups 1st lycopod leave
1st evidence of land plants
460

Mature vessel elements
in Zea mays; open tracheids
perforation plate allows
vessel to act as a water
pipe. SEM. Copyright
John N. A. Lott/BPS.
Sources: Freeman (2002), Purves et al (2001)
A primer on terminology
associated with vascular tissue
Vascular tissue; tissue involved in
transport of materials (water,
minerals…) in plants -- we’re focusing
on water transport in this discussion
Tracheids specialized cells for water
transport; first “true” vascular cells.
Tracheophytes are plants with
Tracheids
Mosses have cells that have a vascular
function, hence its not entirely correct to
refer to them as “nonvascular”, but they
are definitely “non-tracheophyes”, as
are liverworts and hornworts
Tracheophytes have two vascular tissue
types, each composed of several cell
types
xylem; transports water and
dissolved minerals; cell types are
tracheids and, in flowering
Vessel plants, vessel elements
elements phloem; transports sugars, ie,
products of photosynthesis;
minerals too; sieve-tube
members and companion cells
A primer on reproductive cells, embryos, seeds, and
life cycles
Alternation of generations occurs in all plant life cycles,
but not all Charorphyceans; this trait evolved
independently as a derived characteristic of land plants
In some algae with alternation of generations,
sporophytes and gametophytes look similar; not so in
land plants – in all species sporophytes and
gametophytes are very different morphologically
I

Marchantia, a common liverwort. The sporophytes are born within the

tissues of the umbrella shaped structures that arise from the surface
of the flat, green, creeping gametophyte
 A primer on reproductive cells, embryos, seeds, and life
cycles
spores are 1n cells that can divide mitotically, with no syngamy,
to produce individuals
gametes are 1n cells that undergo syngamy to produce a (2n)
multicellular individual
embryo embryonic plant that develop from zygotes within
tissues of the female parent (which provides nutrients)
seed Unique structure in “seed plants”; key structure in
domination of terrestrial habitats by plants; plant embryo and its
food supply, packaged in a protective coat. gymosperms
(conifers and their relatives) produce “naked” seeds in that they
are not born in protective chambers. angiosperms produce
seeds

alternation of Embryo of Marchantia, Embryo of shepherd’s

generations a liverwort purse an angiosperm
Sources: Freeman (2002), Purves et al (2001)

Three ways of expressing (the

same!!) phylogenetic
relationships among major
plant lineages
Cladogram to the left is
particularly useful in that it shows:
-extinct non-seed tracheophyte
lineage Rhyniopsida
-origin of important innovations,
including independent origins --
homoplasies
 Rhynie in the Grampian
Region of Scotland has
become famous as one of
the most important
palaeobotanical localities
in the world. In 1912 the
Scottish geologist
William Mackie
discovered an occurrence
of Lower Devonian
plant-bearing cherts near
this small village.
Recent radiometric
datings of these rocks
give an age of396 ± 12
million years
Rhynie Chert cut perpendicularly to the
stratification showing successive
horizons

Nature 389, 33 - 39 (1997) © Macmillan Publishers Ltd.

The origin and early evolution of plants on land
PAUL KENRICK AND PETER R. CRANE
the Swedish Museum of Natural History, Box 50007, S-10405, Stockholm, SwedenThe Field Museum,
Roosevelt Road at Lake Shore Drive,
Chicago, Illinois60605, Department of the Geophysical Sciences, University of Chicago, USA.
The origin and early evolution of land plants in the mid-Palaeozoic era, between about 480 and 360 million
years ago, was an important event in the history of life, with far-reaching consequences for the evolution of
terrestrial organisms and global environments. A recent surge of interest, catalysed by palaeobotanical
discoveries and advances in the systematics of living plants, provides a revised perspective on the
evolution of early land plants and suggests new directions for future research.

Rubisco catalyzes a reaction in the Calvin-
Benson cycle of photosynthesis in which
carbon from CO2 is added to a five-carbon
chain.
Our current understanding of Plant
phylogenetic relationships is based on
analysis of both morphological and
molecular characters
•SSU rRNA and Rubisco
•presence/absence of vascular tissue,
leaves, seeds…
The broad picture of plant
macroevolutionary patterns includes…
•Divergenge of entire clade from green algae
•All Plantae lineages; cellulose-based cell
walls, chlorophyll a and b, starch as storage
molecule in chloroplasts
•Two of the three earliest lineages (non-
tracheophytes) lack water transport cells;
mosses have have limited numbers of them
•Seedless vascular plants (nonseed
tracheophytes) have vascular tissue and
leaves, but reproduce by making spores; no
seeds
Gymnosperms and Angiosperms have
vascular tissue, they have complex leaves,
and they produce seeds.
Early lineages dependent on wet habitats
and more recent ones not – adaptive
radiations into mesic and xeric conditions
 Dense matt of mosses colonizes old lava in
Iceland. Copyright BPS

Leafy liverwort (Lophocolea bidentata),

showing overlapping leaves
andcreeping growth form. Copyright
BPS.

Leafy liverwort (Lophocolea bidentata),

Marchantia, a common liverwort. The
sporophytes are born within the tissues of
the umbrella shaped structures that arise
from the surface of the flat, green,
creeping gametophyte
Hornwort sporophytes. Unlike
other Bryophytes, hornwort
sporophytes are photosynthetic
showing overlapping leaves and creeping
growth form. Copyright BPS.
Gametophytes of Sphagnum moss growing in a bog in central
Newfoundland. Copyright BPS.
 Order Lycopodiales: Club Moss Big leaf
club moss maples hung with
(Lycopodium epiphytes, most of
annotinum); Denali which are club
National Park and mosses; Hoh Rain
Preserve, AK. Forest, WA. Copyright
Copyright BPS. BPS.

Fern (Blechnum magellanicum) Tree ferns (Dicksonia

in rain forest of Chile's antarctica), in a forest of
Patagonian coast. Copyright Eucalyptus regnans;
Alejandro Frid/BPS. Australia. Copyright
BPS.
Simple determinate
inflorescence of shooting star
(Dodecatheon meadia).
Copyright J. Robert
Stottlemyer/BPS.
Family Arecaceae: desert Family Pinaceae: Colorado
fan palm (Washingtonia Family Pinaceae: Pinus
blue spruce (Picea
filifera); Colorado Desert, ponderosa, ponderosa pine,
pungens). Copyright
CA. Copyright Jon Mark is widespread in the
Pollock/BPS.
Stewart/BPS. American West. Copyright
 The Transition to Land involved evolution of three Sources: Freeman (2002), Purves et al (2001)
20
First extensive important steps
grassland
Preventing cell dessication
Transporting water from cells in plant body with direct
access to water, to cells without
Transporting gametes by a mechanism other than
floating or swimming in water
140
First Flowering
Plant

First nectar-drinking
insects

First vessels
270
(lycopods, seed ferns,
Carboniferous

and horsetails

fossilized spores in tetrads

abundant)

1st seeds The earliest interval of plant

360
1st vascular plant leaves evolution on our time scale
1st wood
1st roots
has yielded fosils of:
1st vascular tissue -spores surrounded by tough
410 1st stomata
1st megafossils (small
membrane made of
club mosses and other sporopollenin
extinct groups and 1st -sheets of a waxy material –
lycopod leave cuticle
460 First evidence of plants -small tubes Fragment of plant cuticle
 Sources: Freeman (2002), Purves et al (2001)
20
First extensive
grassland

140
First Flowering
Plant

Leaf epiderm of dayflower

(Commelina sp.), a monocot. LM.
Copyright Alfred Owczarzak/BPS.
First nectar-drinking
insects Cross-section of a modern leaf (eudicot)
First vessels
270
(lycopods, seed ferns,

Waxy cuticle solved one problem

Carboniferous

and horsetails
abundant)

and caused another – it impeded

uptake of carbon dioxide
modern plants have specialized cells
1st seeds
that solve this problem; two specialized
360 guard cells form a stoma (pore) that
1st vascular plant leaves
1st wood can be open or closed.
1st roots
1st vascular tissue Stomata occur in some extant basal
410 1st stomata lineages -- hornworts and mosses Stomata in leaf of corn, a monocot.
1st megafossils Copyright John N. A. Lott/BPS
(small club mosses Stomata are known from 385 million
and other extinct years ago, from a fossil Rhyniopsid
groups
460 1st lycopod leave
 20
First extensive
grassland

140
First Flowering
Plant

First nectar-drinking
insects

First vessels
270
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

1st seeds
360
1st vascular plant leaves
1st wood
1st roots
1st vascular tissue
410 1st stomata Stomata in leaf of corn, a monocot.
1st megafossils Copyright John N. A. Lott/BPS
(small club mosses
and other extinct Sources: Freeman (2002), Purves et al (2001)
groups
Early Devonian rhyniophyte fossils. f. Stomate with
460 1st lycopod leave tworeniform guard cells (stippled

of structural support and internal water transport
Dense matt of mosses colonizes old lava in
Iceland may be remniscent of early plant life
forms – low and sprawling. Copyright BPS
Artist’s rendering of Carboniferous forest in a
tropical river delta. Most of the plants
depicted here were nonseed tracheophytes
10 to 20 meters tall. In the distance, earl
seed plants up to 40 meters tall towered over
the forest.
Key evolutionary innovations solved the problems
First land plants probably were low, sprawling
before evolution of rigid body support, algal-like tissue
could not grow erect, resist gravity
Before evolution of internal water transport system,
virtually entire plant had to communicate directly with
water
Among communities of low, sprawling plant – growing
horizontally in a sense, competition for space and light
was probably intense –intense selection for erect growth
forms
Transition to terrestrial life required
-internal water transport to high and dry parts of
plant, against the force of gravity
-rigidity of body in order to resist forces of gravity
and wind
 20
First extensive
grassland

140 Cross section of

First Flowering
Plant vascular bundles from a Water-conducting xylem
eudicot stem; nutrient- vessels in a cucumber. LM.
conducting phloem, Copyright J. Robert
water-conducting xylem. Waaland/BPS.
LM. Copyright James
First nectar-drinking Solliday/BPS.
insects

First vessels
270
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

1st seeds Mature vessel elements

360
1st vascular plant leaves in Zea mays; open
1st wood perforation plate allows
1st roots vessel to act as a water
1st vascular tissue
pipe. SEM. Copyright
410 1st stomata
1st megafossils John N. A. Lott/BPS.
(small club mosses Modern day tracheids (left) and
and other extinct vessel elements (right)
groups
460 1st lycopod leave Sources: Freeman (2002), Purves et al (2001)
 Sources: Freeman (2002), Purves et al (2001)
tracheids
20
First extensive
grassland Water transport and structural
pits support
Rhynie chert (~400 mya) bears fossil
plants in an erect, upright postion –
fossils suggest many or most plants
grew erect
140
First Flowering
Kenrick and Crane identifed four types
Plant of water-conducting cells in the Rhynie
vessels
chert (not tracheids or vessel elements)
– some with lignified rings that may
have allowed plants to assume an
upright life form (lignin; strong,
First nectar-drinking lightweight 6-carbon polymer
insects
By about 380 mya, fossil record bears
First vessels tracheids; advanced water conducting
above water- cells found in all modern phyla of
280
(lycopods, seed ferns,

conducting vascular plants. Later tracheids with

Carboniferous

and horsetails

cells from extensive lignification and thickened

abundant)

top; picture of some of the rhynie chert secondary walls

first tracheids 380 mya
from the fossil record; Wood develops from “secondary
diagram shows tracheids xylem”, and arose independently in
1st seeds from modern plant several lineages by about 380 mya.
360
1st vascular plant leaves
bottom; photo of 50 my old First trees appear in fossil record some
1st wood
1st roots vessels and diagram of 370 mya
1st vascular tissue modern vessels. Vessels
410 1st stomata are shorter and fatter than
1st megafossils tracheids, and stacked end
(small club mosses
to end. Water flows up
and other extinct
groups vessels directly, and
laterally, from vessel to
460 1st lycopod leave
vessl, as well, through
perforations in primary and
Evolution of heterospory
and male and female
gametophytes
The earliest tracheophytes
were homosporous.
Heterospory evolved multiple
times in tracheophytes in
early evolution of land plants
(occurs in Rhyniophyte
lineage)
Multiple evolutionary origins
suggests heterospory affords
selective advantages;
Later tracheophytes,
especially seed plants, show
ever greater specialiation of
the heterosporous condition Homosporous plants have a single type of Heterosporous plants have two types
spore that gives rise to a single type of of spores, each of which develops into
gametophyte, which bears two types of sex male or female gametophytes, each of
organs which produces eggs or sperm
 20 Protecting and transporting Embryos: Evolution of seed and pollination
First extensive
grassland
All land plants have life cycles with alternation of generations.
Relationship between gametophyte and sporophyte varies among lineages in terms
of size, dependence, dominance in life cycle
Mosses: sporophyte is small and depends on gametophyte for nutrition
Ferns; sporophyte is larger than gametophyte and is autotrophic
140 (photosynthesizes its own food)
First Flowering
Plant Seed plants (gymnosperms and angiosperms); sporophyte is dominant;
-male gametophyte is reduced to pollen grain
-female gametophyte to a small structure that holds the egg or eggs

First nectar-drinking
insects

First vessels
270
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

1st seeds
360
1st vascular plant leaves
1st wood
1st roots
1st vascular tissue
410 1st stomata
1st megafossils (small
club mosses and other Gametophytes (green) of the star moss
extinct groups and 1st produce non-photosynthetic stalked
lycopod leaves
First evidence of
sporophytes with caplike sporangia at their
460 tips (Campbell 2002)
land plants
 20 Protecting and transporting Embryos: Evolution of pollen, pollination and seed and seed
First extensive dispersal
grassland
Seed plants (gymnosperms and angiosperms); sporophyte is dominant;
-male gametophyte is reduced to pollen grain
-female gametophyte to a small structure that holds the egg or eggs
Selection favored male gametophytes highly reduced in size and encased in
sporopollenin; pollen grains are the consequence of that bout of mutation and
140
First Flowering selection
Plant
Pollen grains survive for prolonged periods in dry environments without
becoming dessicated
Pollen grains can be transported by wind, gravity or animals; through pollen
grains, seed plants lost dependence on water for fertilization
First nectar-drinking
insects

First vessels
270
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

1st seeds
360
1st vascular plant leaves
1st wood
1st roots Pollination. Carrion plant (left) has foul odor that attracts carrion flies.
1st vascular tissue Hummingbird pollinated flower (middle) produces nectar. Many arge,
410 1st stomata complex flowers (eg sunflower, right) attract a host of species that act
1st megafossils (small as pollinators
club mosses and other
extinct groups and 1st
lycopod leaves
First evidence of
460
land plants
 20 Evolution of seeds and flowers
First extensive
grassland
Seeds enclose and protects embryo, and has a stockpile of nutrients
Seeds are often attached to a structure that facilitates dispersal by wind, water or
animals

dispersal by wind dispersal by water

140
First Flowering
Plant

First nectar-drinking
insects

First vessels
270
(lycopods, seed ferns,

dispersal by animals
Carboniferous

and horsetails
abundant)

1st seeds
360
1st vascular plant leaves
1st wood
1st roots
1st vascular tissue
410 1st stomata
1st megafossils (small
club mosses and other
extinct groups and 1st
lycopod leaves
First evidence of
460
land plants
Extra slides
 2 Sources: Freeman (2002), Purves et al (2001)
20
First extensive grassland

Overview of the evolutionary history of plants

Consider five intervals of evolutionary history, each defined
by at least one major evolutionary innovation
Archaefructus, an early flower

140
First Flowering Plant

Cones from Araucaria mirabilis, an early gymnsperm

First nectar-drinking insects

Fossil frond of fern (Asterotheca
arborescens); late Carboniferous.
First vessels
Copyright Barbara J. Miller/BPS.
270
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

Fossil stem of oldest known

lycopod genus (Bothrodendron
minutifolium); late Devonian. LM.
Seed fern leaves
Copyright Phil Gates/BPS.
360 1st seeds
1st vascular plant leaves Cells in fossil stem of Rhynia
1st wood major (extinct Phylum
1st roots Rhyniophyta); late Silurian and
1st vascular tissue
1st stomata
Devonian. LM. Copyright Phil
410
Gates/BPS.
1st megafossils
(small club
Spores in fossil Rhynia major; late
mosses and
other extinct Silurian and Devonian. LM.
groups
Fragment of plant cuticle
Copyright Phil Gates/BPS
1st lycopod
460 leave
 20
First extensive
grassland

140
First Flowering
Plant

First nectar-drinking
insects

First vessels
270
(lycopods, seed ferns,
Carboniferous

and horsetails
abundant)

1st seeds
360
1st vascular plant leaves
1st wood
1st roots
1st vascular tissue
410 1st stomata
1st megafossils (small
club mosses and other
extinct groups and 1st
lycopod leaves
First evidence of
460
land plants
Key Evolutionary Innovations and Trends in the
Transition of Plants to Land
-reducing water loss; cuticle and stomata
-transporting water; vascular tissue and wood
-transporting gametes and prtoecting embryos;
pollination mechanisms and nutrititive, protective
seeds
Key evolutionary innovations and trends for
capturing energy from sunlight
-Photosynthetic Pathways C3 and C4
-Crassulacean acid metabolism
-growth habits and life forms
Minimizing water loss and regulating gas exchange
Nature 389, 33 - 39 (1997) © Macmillan Publishers Ltd.
The origin and early evolution of plants on land
PAUL KENRICK AND PETER R. CRANE
the Swedish Museum of Natural History, Box 50007, S-10405, Stockholm, SwedenThe Field Museum,
Roosevelt Road at Lake Shore Drive,
Chicago, Illinois60605, Department of the Geophysical Sciences, University of Chicago, USA.

The origin and early evolution of land plants in the mid-Palaeozoic era, between about 480 and 360
million years ago, was an important event in the history of life, with far-reaching consequences for
the evolution of terrestrial organisms and global environments. A recent surge of interest, catalysed
by palaeobotanical discoveries and advances in the systematics of living plants, provides a revised
perspective on the evolution of early land plants and suggests new directions for future research.
Figure 1 Morphological diversity among basal living land plants and potential land-plant sister groups.
a,Coleochaete orbicularis (Charophyceae) gametophyte; magnification × 75 (photograph courtesy of L.
E.Graham). b, Chara (Charophyoceae) gametophyte; magnification × 1.5 (photograph courtesy of M.Feist).
c, Riccia (liverwort) gametophyte showing sporangia (black) embedded in the thallus; magnification× 5
(photograph courtesy of A. N. Drinnan). d, Anthoceros (hornwort) gametophyte showing
nbranchedsporophytes; magnification × 2.5 (photograph courtesy of A. N. Drinnan). e, Mnium (moss)
gametophyteshowing unbranched sporophytes with terminal sporangia (capsule); magnification × 4.5
(photographcourtesy of W. Burger). f, Huperzia (clubmoss) sporophyte with leaves showing sessile yellow
sporangia;magnification × 0.8. g, Dicranopteris (fern) sporophyte showing leaves with circinate
vernation;magnification × 0.08. h, Psilotum (whisk fern) sporophyte with reduced leaves and spherical
synangia(three fused sporangia); magnification × 0.4. i, Equisetum (horsetail) sporophyte with whorled
branches,reduced leaves, and a terminal cone; magnification × 0.4. j, Cycas (seed plant) sporophyte
showing leavesand terminal cone with seeds; magnification × 0.05 (photograph courtesy of W. Burger).
Figure 2 a, Longitudinal section of part of a silicified early fossil gametophyte (Kidstonophyton discoides
from the Rhynie Chert). Antheridia (male sexual organs) are located on the upper surface of the branch;
magnification × 3.4. b, Longitudinal section of antheridium of Lyonophyton rhyniensis from the Rhynie
Chert; magnification × 40. c, Longitudinal section of archegonium (female sexual organ) of Langiophyton
mackiei from the Rhynie Chert; magnification × 80. a–c are from the Remy Collection (slides 200, 90 and
330), Abteilung Paläobotanik, Westfälische Wilhelms-Universität, Münster, Germany (photographs
courtesy of H. Hass and H. Kerp). d, Scanning electron micrograph of Tetrahedraletes medinensis
showing a spore tetrad of possible liverwort affinity from the Late Ordovician (photograph courtesy of W.
A. Taylor); magnification × 670.
Figure 3 Sporophyte diversity in Early Devonian rhyniophyte fossils. a, Cooksonia pertoniiapiculispora:
sporophyte (incomplete proximally) with terminal sporangium15; magnification × 15. b,Tortilicaulis offaeus:
sporophyte (incomplete proximally) with terminal sporangium81; magnification × 40.c. Tortilicaulis offaeus:
sporophyte (incomplete proximally) with terminal bifurcating sporangium81;magnification × 30. d, Transverse
section of sporangium showing thick wall and central spore mass;magnification × 70. e, Details of epidermis at
rim of sporangium; magnification × 45. f, Stomate with tworeniform guard cells (stippled); magnification × 120.
Figure 5 Diversity of water-conducting cells (tracheids) in early land plants (median
longitudinal sectionthrough cells, basal and proximal end wa. lls not shown; cells are 20–40
m diameter). a, Top,bryophyte hydroid; bottom, details of hydroid wall showing distribution of
plasmodesmata-derivedmicropores (10–50 nm diameter; stipple)84. b, Top, S-type tracheid
(fossil) of Rhyniopsida; bottom,details of S-type cell wall showing distribution of
plasmodesmata-derived micropores (stipple) and'spongy' interior to thickenings19. c, Top,
G-type tracheid (fossil) of basal extinct eutracheophytes, whichclosely resemble the
tracheids of some living vascular plants; bottom, details of G-type cell wall showingpores
distributed between thickenings19. d, Top, scalariform pitted P-type tracheid (fossil) typical
oftrimerophyte grade plants (euphyllophytes); bottom, details of P-type cell wall showing pit
chambers andsheet with pores that extends over pit apertures26.