Embryophyte

The Embryophyta (/ɛmbriˈɒfɪtə,

ˈɛmbrioʊfaɪtə/) or land plants are the
most familiar group of green plants that
form vegetation on earth. Embryophyta is
a clade within the Phragmoplastophyta, a
larger clade that also includes several
green algae groups (including the
Charophyceae and Coleochaetales), and
within this large clade the embryophytes
(/ˈɛmbriəfaɪts/) are sister to the
Zygnematophyceae/Mesotaeniaceae
and consist of the bryophytes plus the
polysporangiophytes.[12] Living
embryophytes therefore include
hornworts, liverworts, mosses, ferns,
lycophytes, gymnosperms and flowering
plants.
 Land plants
Temporal range: Mid Ordovician–Present[1][2]
PreꞒ Ꞓ O S D C P T J K PN
g
(Spores from Dapingian (early Middle
Ordovician))

Scientific classification

Kingdom: Plantae

Clade: Streptophyta

Clade: Embryophytes
Engler, 1892[3][4]
 Divisions
Non-vascular land plants
(bryophytes)
Marchantiophyta – liverworts
Bryophyta – mosses
Anthocerotophyta – hornworts
†Horneophytopsida
Vascular plants (tracheophytes)
†Rhyniophyta – rhyniophytes
†Zosterophyllophyta –
zosterophylls
Lycopodiophyta – clubmosses
†Trimerophytophyta –
trimerophytes
Polypodiophyta – ferns and
horsetails
 Seed plants (spermatophytes)
†Pteridospermatophyta –
seed ferns
Pinophyta – conifers
Cycadophyta – cycads
Ginkgophyta – ginkgo
Gnetophyta – gnetae
Magnoliophyta – flowering
plants

Traditional groups:

Bryophyta*
Pteridophyta*
Gymnospermae*
Angiospermae
 Synonyms
Cormophyta Endlicher, 1836
Phyta Barkley, 1939[5]
Cormobionta Rothmaler, 1948[6]
Euplanta Barkley, 1949[7]
Telomobionta Takhtajan, 1964[8]
Embryobionta Cronquist et al. 1966[9]
Metaphyta Whittaker, 1969[10]
Plantae Margulis, 1971[11]

The Embryophyta are informally called

land plants because they live primarily in
terrestrial habitats, while the related
green algae are primarily aquatic.
Embryophytes are complex multicellular
eukaryotes with specialized reproductive
organs. The name derives from their
innovative characteristic of nurturing the
young embryo sporophyte during the
early stages of its multicellular
development within the tissues of the
parent gametophyte. With very few
exceptions, embryophytes obtain their
energy by photosynthesis, that is by
using the energy of sunlight to synthesize
their food from carbon dioxide and water.

Description
Moss, clubmoss, ferns and cycads in a greenhouse

The evolutionary origins of the

embryophytes are discussed further
below, but they are believed to have
evolved from within a group of complex
green algae during the Paleozoic era
(which started around
540 million years ago)[13][14] probably
from terrestrial unicellular charophytes,
similar to extant
Klebsormidiophyceae.[15] Embryophytes
are primarily adapted for life on land,
although some are secondarily aquatic.
Accordingly, they are often called land
plants or terrestrial plants.

On a microscopic level, the cells of

embryophytes are broadly similar to
those of green algae, but differ in that in
cell division the daughter nuclei are
separated by a phragmoplast.[16] They
are eukaryotic, with a cell wall composed
of cellulose and plastids surrounded by
two membranes. The latter include
chloroplasts, which conduct
photosynthesis and store food in the
form of starch, and are characteristically
pigmented with chlorophylls a and b,
generally giving them a bright green
color. Embryophyte cells also generally
have an enlarged central vacuole
enclosed by a vacuolar membrane or
tonoplast, which maintains cell turgor
and keeps the plant rigid.

In common with all groups of

multicellular algae they have a life cycle
which involves 'alternation of
generations'. A multicellular generation
with a single set of chromosomes – the
haploid gametophyte – produces sperm
and eggs which fuse and grow into a
multicellular generation with twice the
number of chromosomes – the diploid
sporophyte. The mature sporophyte
produces haploid spores which grow into
a gametophyte, thus completing the
cycle. Embryophytes have two features
related to their reproductive cycles which
distinguish them from all other plant
lineages. Firstly, their gametophytes
produce sperm and eggs in multicellular
structures (called 'antheridia' and
'archegonia'), and fertilization of the
ovum takes place within the
archegonium rather than in the external
environment. Secondly, and most
importantly, the initial stage of
development of the fertilized egg (the
zygote) into a diploid multicellular
sporophyte, take place within the
archegonium where it is both protected
and provided with nutrition. This second
feature is the origin of the term
'embryophyte' – the fertilized egg
develops into a protected embryo, rather
than dispersing as a single cell.[13] In the
bryophytes the sporophyte remains
dependent on the gametophyte, while in
all other embryophytes the sporophyte
generation is dominant and capable of
independent existence.

Embryophytes also differ from algae by

having metamers. Metamers are
repeated units of development, in which
each unit derives from a single cell, but
the resulting product tissue or part is
largely the same for each cell. The whole
organism is thus constructed from
similar, repeating parts or metamers.
Accordingly, these plants are sometimes
termed 'metaphytes' and classified as the
group Metaphyta[17] (but Haeckel's
definition of Metaphyta places some
algae in this group[18]). In all land plants a
disc-like structure called a phragmoplast
forms where the cell will divide, a trait
only found in the land plants in the
streptophyte lineage, some species
within their relatives Coleochaetales,
Charales and Zygnematales, as well as
within subaerial species of the algae
order Trentepohliales, and appears to be
essential in the adaptation towards a
terrestrial life style.[19][20][21][22]

Phylogeny and classification

All green algae and land plants are now
known to form a single evolutionary
lineage or clade, one name for which is
Viridiplantae (i.e. 'green plants').
According to several molecular clock
estimates the Viridiplantae split
1,200 million years ago to
725 million years ago into two clades:
chlorophytes and streptophytes. The
chlorophytes are considerably more
diverse (with around 700 genera) and
were originally marine, although some
groups have since spread into fresh
water. The streptophyte algae (i.e. the
streptophyte clade minus the land
plants) are less diverse (with around 122
genera) and adapted to fresh water very
early in their evolutionary history. They
have not spread into marine
environments (only a few stoneworts,
which belong to this group, tolerate
brackish water). Some time during the
Ordovician period (which started around
490 million years ago) one or more
streptophytes invaded the land and
began the evolution of the embryophyte
land plants.[23] Present day
embryophytes form a monophyletic
group called the hemitracheophytes.[24]
Becker and Marin speculate that land
plants evolved from streptophytes rather
than any other group of algae because
streptophytes were adapted to living in
fresh water. This prepared them to
tolerate a range of environmental
conditions found on land. Fresh water
living made them tolerant of exposure to
rain; living in shallow pools required
tolerance to temperature variation, high
levels of ultra-violet light and seasonal
dehydration.[25]

Relationships between the groups

making up Viridiplantae are still being
elucidated. Views have changed
considerably since 2000 and
classifications have not yet caught up.
However, the division between
chlorophytes and streptophytes and the
evolution of embryophytes from within
the latter group, as shown in the
cladogram below, are well
established.[23][26] Three approaches to
classification are shown. Older
classifications, as on the left, treated all
green algae as a single division of the
plant kingdom under the name
Chlorophyta.[27] Land plants were then
placed in separate divisions. All the
streptophyte algae can be grouped into
one paraphyletic taxon, as in the middle,
allowing the embryophytes to form a
taxon at the same level. Alternatively, the
embryophytes can be sunk into a
monophyletic taxon comprising all the
streptophytes, as shown below.[26] A
variety of names have been used for the
different groups which result from these
approaches; those used below are only
one of a number of possibilities. The
higher-level classification of the
Viridiplantae varies considerably,
resulting in widely different ranks being
assigned to the embryophytes, from
kingdom to class.
   chlorophytes
 
streptophyte
Viridiplantae   algae
  streptophytes   (paraphyletic
group)
 
  embryophyte
 

The precise relationships within the

streptophytes are less clear as of
March 2012. The stoneworts (Charales)
have traditionally been identified as
closest to the embryophytes, but recent
work suggests that either the
Zygnematales or a clade consisting of
the Zygnematales and the
Coleochaetales may be the sister group
to the land plants.[28][29] That the
Zygnematales (or Zygnematophyceae)
are the closest algal relatives to land
plants was underpinned by an exhaustive
phylogenetic analysis (phylogenomics)
performed in 2014,[30] which is supported
by both plastid genome phylogenies[31]
as well as plastid gene content and
properties.[32]

The preponderance of molecular

evidence as of 2006 suggested that the
groups making up the embryophytes are
related as shown in the cladogram below
(based on Qiu et al. 2006 with additional
names from Crane et al. 2004).[33][34]
   Liverworts
 
  Mosses
 
  Ho
 

Living embryophytes
   
  
  Tracheophytes
  E

Studies based on morphology rather than

on genes and proteins have regularly
reached different conclusions; for
example that neither the monilophytes
(ferns and horsetails) nor the
gymnosperms are a natural or
monophyletic group.[35][36][37]

There is considerable variation in how

these relationships are converted into a
formal classification. Consider the
angiosperms or flowering plants. Many
botanists, following Lindley in 1830, have
treated the angiosperms as a division.[38]
Palaeobotanists have usually followed
Banks in treating the tracheophytes or
vascular plants as a division,[39] so that
the angiosperms become a class or even
a subclass. Two very different systems
are shown below. The classification on
the left is a traditional one, in which ten
living groups are treated as separate
divisions; the classification on the right
(based on Kenrick and Crane's 1997
treatment) sharply reduces the rank of
groups such as the flowering plants.[40]
(More complex classifications are
needed if extinct plants are included.)
Two contrasting classifications of living
land plants
Liverworts Marchiantiophyta Marchiant
Mosses Bryophyta Bryophyta
Hornworts Anthocerotophyta Anthocero
Tracheop
Lycophytes Lycopodiophyta Lycophyt
Euphyllo
Ferns and
Pteridophyta Monilifo
horsetails
Radiato
Cycads Cycadophyta Cycada
Conifers Pinophyta Conife
Ginkgo Ginkgophyta Ginkgo
Gnetophytes Gnetophyta Anthop
Flowering Magnoliophyta
plants

An updated phylogeny of Embryophytes

based on the work by Novíkov & Barabaš-
Krasni 2015[41] and Hao and Xue 2013[42]
with plant taxon authors from Anderson,
Anderson & Cleal 2007[43] and some
clade names from Pelletier 2012 and
others.[44][45] Puttick et al./Nishiyama et
al are used for the basal clades.[12][46][47]
Embryophytes   Ant
   
Bryomorpha  B
  Setaphyta  
   M
 
Polysporangiomorpha  
   
Tracheophytin
 
Diversity

Bryophytes …

Most bryophytes, such as these mosses, produce

stalked sporophytes from which their spores are
released.

Bryophytes consist of all non-vascular

land plants (embryophytes without
vascular tissue). All are relatively small
and are usually confined to environments
that are humid or at least seasonally
moist. They are limited by their reliance
on water needed to disperse their
gametes, although only a few bryophytes
are truly aquatic. Most species are
tropical, but there are many arctic
species as well. They may locally
dominate the ground cover in tundra and
Arctic–alpine habitats or the epiphyte
flora in rain forest habitats.

The three living divisions are the mosses

(Bryophyta), hornworts
(Anthocerotophyta), and liverworts
(Marchantiophyta). Originally, these three
groups were included together as
classes within the single division
Bryophyta. They have usually been
placed separately into three divisions
under the assumption that the
bryophytes are a paraphyletic (more than
one lineage) group, but newer research
supports the monophyletic (having a
common ancestor) model.[48] The three
bryophyte groups form an evolutionary
grade of those land plants that are not
vascular. Some closely related green
algae are also non-vascular, but are not
considered "land plants".

Marchantiophyta (liverworts)
Bryophyta (mosses)
Anthocerotophyta (hornworts)

Regardless of their evolutionary origins,

the bryophytes are usually studied
together because of their many
biological similarities as non-vascular
land plants. All three bryophyte groups
share a haploid-dominant (gametophyte)
life cycle and unbranched sporophytes
(the plant's diploid structure). These are
traits that appear to be plesiotypic within
the land plants, and thus were common
to all early diverging lineages of plants on
the land. The fact that the bryophytes
have a life cycle in common may thus be
an artefact of being the oldest extant
lineages of land plant, and not the result
of close shared ancestry. (See the
phylogeny above.)
The bryophyte life-cycle is strongly
dominated by the haploid gametophyte
generation. The sporophyte remains
small and dependent on the parent
gametophyte for its entire brief life. All
other living groups of land plants have a
life cycle dominated by the diploid
sporophyte generation. It is in the diploid
sporophyte that vascular tissue
develops. Although some mosses have
quite complex water-conducting vessels,
bryophytes lack true vascular tissue.

Like the vascular plants, bryophytes do

have differentiated stems, and although
these are most often no more than a few
centimeters tall, they do provide
mechanical support. Most bryophytes
also have leaves, although these typically
are one cell thick and lack veins. Unlike
the vascular plants, bryophytes lack true
roots or any deep anchoring structures.
Some species do grow a filamentous
network of horizontal stems, but these
have a primary function of mechanical
attachment rather than extraction of soil
nutrients (Palaeos 2008).

Rise of vascular plants …

Reconstruction of a plant of Rhynia

During the Silurian and Devonian periods

(around 440 to 360 million years ago),
plants evolved which possessed true
vascular tissue, including cells with walls
strengthened by lignin (tracheids). Some
extinct early plants appear to be between
the grade of organization of bryophytes
and that of true vascular plants
(eutracheophytes). Genera such as
Horneophyton have water-conducting
tissue more like that of mosses, but a
different life-cycle in which the
sporophyte is more developed than the
gametophyte. Genera such as Rhynia
have a similar life-cycle but have simple
tracheids and so are a kind of vascular
plant. It was assumed that the
gametophyte dominant phase seen in
bryophytes used to be the ancestral
condition in terrestrial plants, and that
the sporophyte dominant stage in
vascular plants was a derived trait. But
research point out the possibility that
both the gametophyte and sporophyte
stage were equally independent from
each other, and that the mosses and
vascular plants in that case are both
derived, and has evolved in the opposite
direction from the other.[49]

During the Devonian period, vascular

plants diversified and spread to many
different land environments. In addition
to vascular tissues which transport water
throughout the body, tracheophytes have
an outer layer or cuticle that resists
drying out. The sporophyte is the
dominant generation, and in modern
species develops leaves, stems and
roots, while the gametophyte remains
very small.

Lycophytes and euphyllophytes …

Lycopodiella inundata, a lycophyte

All the vascular plants which disperse
through spores were once thought to be
related (and were often grouped as 'ferns
and allies'). However, recent research
suggests that leaves evolved quite
separately in two different lineages. The
lycophytes or lycopodiophytes – modern
clubmosses, spikemosses and quillworts
– make up less than 1% of living vascular
plants. They have small leaves, often
called 'microphylls' or 'lycophylls', which
are borne all along the stems in the
clubmosses and spikemosses, and
which effectively grow from the base, via
an intercalary meristem.[50] It is believed
that microphylls evolved from
outgrowths on stems, such as spines,
which later acquired veins (vascular
traces).[51]

Although the living lycophytes are all

relatively small and inconspicuous
plants, more common in the moist
tropics than in temperate regions, during
the Carboniferous period tree-like
lycophytes (such as Lepidodendron)
formed huge forests that dominated the
landscape.[52]

The euphyllophytes, making up more

than 99% of living vascular plant species,
have large 'true' leaves (megaphylls),
which effectively grow from the sides or
the apex, via marginal or apical
meristems.[50] One theory is that
megaphylls developed from three-
dimensional branching systems by first
'planation' – flattening to produce a two
dimensional branched structure – and
then 'webbing' – tissue growing out
between the flattened branches.[53]
Others have questioned whether
megaphylls developed in the same way
in different groups.[54]

Ferns and horsetails …

Athyrium filix-femina, unrolling young frond

Euphyllophytes are divided into two
lineages: the ferns and horsetails
(monilophytes) and the seed plants
(spermatophytes). Like all the preceding
groups, the monilophytes continue to use
spores as their main method of
dispersal. Traditionally, whisk ferns and
horsetails were treated as distinct from
'true' ferns. Recent research suggests
that they all belong together,[55] although
there are differences of opinion on the
exact classification to be used. Living
whisk ferns and horsetails do not have
the large leaves (megaphylls) which
would be expected of euphyllophytes.
However, this has probably resulted from
reduction, as evidenced by early fossil
horsetails, in which the leaves are broad
with branching veins.[56]

Ferns are a large and diverse group, with

some 12,000 species.[57] A stereotypical
fern has broad, much divided leaves,
which grow by unrolling.

Seed plants …
Pine forest in France

Large seed of a horse chestnut, Aesculus

hippocastanum

Seed plants, which first appeared in the

fossil record towards the end of the
Paleozoic era, reproduce using
desiccation-resistant capsules called
seeds. Starting from a plant which
disperses by spores, highly complex
changes are needed to produce seeds.
The sporophyte has two kinds of spore-
forming organs (sporangia). One kind,
the megasporangium, produces only a
single large spore (a megaspore). This
sporangium is surrounded by one or
more sheathing layers (integuments)
which form the seed coat. Within the
seed coat, the megaspore develops into
a tiny gametophyte, which in turn
produces one or more egg cells. Before
fertilization, the sporangium and its
contents plus its coat is called an 'ovule';
after fertilization a 'seed'. In parallel to
these developments, the other kind of
sporangium, the microsporangium,
produces microspores. A tiny
gametophyte develops inside the wall of
a microspore, producing a pollen grain.
Pollen grains can be physically
transferred between plants by the wind or
animals, most commonly insects. Pollen
grains can also transfer to an ovule of the
same plant, either with the same flower
or between two flowers of the same
plant (self-fertilization). When a pollen
grain reaches an ovule, it enters via a
microscopic gap in the coat (the
micropyle). The tiny gametophyte inside
the pollen grain then produces sperm
cells which move to the egg cell and
fertilize it.[58] Seed plants include two
groups with living members, the
gymnosperms and the angiosperms or
flowering plants. In gymnosperms, the
ovules or seeds are not further enclosed.
In angiosperms, they are enclosed in
ovaries. A split ovary with a visible seed
can be seen in the adjacent image.
Angiosperms typically also have other,
secondary structures, such as petals,
which together form a flower.

Extant seed plants are divided into five

groups:

Gymnosperms
Pinophyta - conifers
Cycadophyta - cycads
Ginkgophyta - ginkgo
 Gnetophyta - gnetophytes
Angiosperms
Magnoliophyta – flowering plants

References
1. Gray, J.; Chaloner, W.G. & Westoll,
T.S. (1985), "The Microfossil Record
of Early Land Plants: Advances in
Understanding of Early
Terrestrialization, 1970-1984 [and
Discussion]", Philosophical
Transactions of the Royal Society B:
Biological Sciences, 309 (1138):
167–195,
Bibcode:1985RSPTB.309..167G ,
doi:10.1098/rstb.1985.0077
2. Rubinstein, C.V.; Gerrienne, P.; De La
Puente, G.S.; Astini, R.A. & Steemans,
P. (2010), "Early Middle Ordovician
evidence for land plants in Argentina
(eastern Gondwana)", New
Phytologist, 188 (2): 365–9,
doi:10.1111/j.1469-
8137.2010.03433.x ,
PMID 20731783
3. Engler, A. 1892. Syllabus der
Vorlesungen über specielle und
medicinisch-pharmaceutische
Botanik: Eine Uebersicht über das
ganze Pflanzensystem mit
Berücksichtigung der Medicinal- und
Nutzpflanzen. Berlin: Gebr.
Borntraeger.
4. Pirani, J. R.; Prado, J. (2012).
"Embryopsida, a new name for the
class of land plants" (PDF). Taxon.
61 (5): 1096–1098.
doi:10.1002/tax.615014 .
5. Barkley, Fred A. Keys to the phyla of
organisms. Missoula, Montana.
1939.
6. Rothmaler, Werner. Über das
natürliche System der Organismen.
Biologisches Zentralblatt. 67: 242-
250. 1948.
7. Barkley, Fred A. "Un esbozo de
clasificación de los organismos" .
Revista de la Facultad Nacional de
 Agronomia, Universidad de
Antioquia, Medellín. 10: 83–103.
8. Takhtajan, A (1964). "The taxa of the
higher plants above the rank of
order" (PDF). Taxon. 13 (5): 160–
164. doi:10.2307/1216134 .
JSTOR 1216134 .
9. Cronquist, A.; Takhtajan, A.;
Zimmermann, W. (1966). "On the
Higher Taxa of Embryobionta"
(PDF). Taxon. 15 (4): 129–134.
doi:10.2307/1217531 .
JSTOR 1217531 .
10. Whittaker, R. H. (1969). "New
concepts of kingdoms or
organisms" (PDF). Science. 163
 (3863): 150–160.
Bibcode:1969Sci...163..150W .
CiteSeerX 10.1.1.403.5430 .
doi:10.1126/science.163.3863.150 .
PMID 5762760 . Archived from the
original (PDF) on 2017-11-17.
Retrieved 2014-11-28.
11. Margulis, L (1971). "Whittaker's five
kingdoms of organisms: minor
revisions suggested by
considerations of the origin of
mitosis". Evolution. 25 (1): 242–245.
doi:10.2307/2406516 .
JSTOR 2406516 . PMID 28562945 .
12. Puttick, Mark N.; Morris, Jennifer L.;
Williams, Tom A.; Cox, Cymon J.;
 Edwards, Dianne; Kenrick, Paul;
Pressel, Silvia; Wellman, Charles H.;
Schneider, Harald (2018). "The
Interrelationships of Land Plants and
the Nature of the Ancestral
Embryophyte" . Current Biology. 28
(5): 733–745.e2.
doi:10.1016/j.cub.2018.01.063 .
PMID 29456145 .
13. Niklas, K.J.; Kutschera, U. (2010),
"The evolution of the land plant life
cycle", New Phytologist, 185 (1): 27–
41, doi:10.1111/j.1469-
8137.2009.03054.x ,
PMID 19863728 .
14. de Vries, J; Archibald, JM (March
2018). "Plant evolution: landmarks
on the path to terrestrial life" . The
New Phytologist. 217 (4): 1428–
1434. doi:10.1111/nph.14975 .
PMID 29318635 .
15. Del-Bem, Luiz-Eduardo (2018-05-31).
"Xyloglucan evolution and the
terrestrialization of green plants" .
New Phytologist. 219 (4): 1150–
1153. doi:10.1111/nph.15191 .
ISSN 0028-646X . PMID 29851097 .
16. Pickett-Heaps, J. (1976). "Cell
division in eucaryotic algae".
BioScience. 26 (7): 445–450.
 doi:10.2307/1297481 .
JSTOR 1297481 .
17. Mayr, E. (1990), "A natural system of
organisms", Nature, 348 (6301): 491,
Bibcode:1990Natur.348..491M ,
doi:10.1038/348491a0 ,
S2CID 13454722
18. Haeckel, Ernst Heinrich Philipp
August (28 September 1894).
"Systematische phylogenie" . Berlin :
Georg Reimer – via Internet Archive.
19. John, Whitfield (19 February 2001).
"Land plants divided and ruled" .
Nature News.
doi:10.1038/conference010222-8 .
20. "Phragmoplastin, green algae and
the evolution of cytokinesis" .
21. "Invasions of the Algae -
ScienceNOW - News - Science" .
Archived from the original on 2013-
06-02. Retrieved 2013-03-27.
22. "All Land Plants Evolved From Single
Type of Algae, Scientists Say" .
23. Becker, B. & Marin, B. (2009),
"Streptophyte algae and the origin of
embryophytes", Annals of Botany,
103 (7): 999–1004,
doi:10.1093/aob/mcp044 ,
PMC 2707909 , PMID 19273476
24. Lecointre, Guillaume; Guyader, Hervé
Le (August 28, 2006). The Tree of
 Life: A Phylogenetic Classification .
Harvard University Press. p. 175 .
ISBN 9780674021839 – via Internet
Archive. "The hemitracheophytes
form a monophyletic group that
unites the bryophytes and the
tracheophytes (or vascular plants)."
25. Becker & Marin 2009, p. 1001
26. Lewis, Louise A. & McCourt, R.M.
(2004), "Green algae and the origin
of land plants", Am. J. Bot., 91 (10):
1535–1556,
doi:10.3732/ajb.91.10.1535 ,
PMID 21652308
27. Taylor, T.N.; Taylor, E.L. & Krings, M.
(2009), Paleobotany, The Biology and
 Evolution of Fossil Plants (2nd ed.),
Amsterdam; Boston: Academic
Press, ISBN 978-0-12-373972-8, p.
1027
28. Wodniok, Sabina; Brinkmann,
Henner; Glöckner, Gernot; Heidel,
Andrew J.; Philippe, Hervé;
Melkonian, Michael & Becker,
Burkhard (2011), "Origin of land
plants: Do conjugating green algae
hold the key?", BMC Evolutionary
Biology, 11 (1): 104,
doi:10.1186/1471-2148-11-104 ,
PMC 3088898 , PMID 21501468
29. Leliaert, Frederik; Verbruggen,
Heroen & Zechman, Frederick W.
 (2011), "Into the deep: New
discoveries at the base of the green
plant phylogeny", BioEssays, 33 (9):
683–692,
doi:10.1002/bies.201100035 ,
PMID 21744372 , S2CID 40459076
30. Wickett, Norman J.; Mirarab, Siavash;
Nguyen, Nam; Warnow, Tandy;
Carpenter, Eric; Matasci, Naim;
Ayyampalayam, Saravanaraj; Barker,
Michael S.; Burleigh, J. Gordon
(2014-11-11). "Phylotranscriptomic
analysis of the origin and early
diversification of land plants" .
Proceedings of the National
Academy of Sciences. 111 (45):
E4859–E4868.
 Bibcode:2014PNAS..111E4859W .
doi:10.1073/pnas.1323926111 .
ISSN 0027-8424 . PMC 4234587 .
PMID 25355905 .
31. Ruhfel, Brad R.; Gitzendanner,
Matthew A.; Soltis, Pamela S.; Soltis,
Douglas E.; Burleigh, J. Gordon
(2014-01-01). "From algae to
angiosperms–inferring the
phylogeny of green plants
(Viridiplantae) from 360 plastid
genomes" . BMC Evolutionary
Biology. 14: 23. doi:10.1186/1471-
2148-14-23 . ISSN 1471-2148 .
PMC 3933183 . PMID 24533922 .
32. Vries, Jan de; Stanton, Amanda;
Archibald, John M.; Gould, Sven B.
(2016-02-16). "Streptophyte
Terrestrialization in Light of Plastid
Evolution". Trends in Plant Science.
21 (6): 467–476.
doi:10.1016/j.tplants.2016.01.021 .
ISSN 1360-1385 . PMID 26895731 .
33. Qiu, Y.L.; Li, L.; Wang, B.; Chen, Z.;
et al. (2006), "The deepest
divergences in land plants inferred
from phylogenomic evidence",
Proceedings of the National
Academy of Sciences, 103 (42):
15511–6,
Bibcode:2006PNAS..10315511Q ,
 doi:10.1073/pnas.0603335103 ,
PMC 1622854 , PMID 17030812
34. Crane, P.R.; Herendeen, P. & Friis,
E.M. (2004), "Fossils and plant
phylogeny" , American Journal of
Botany, 91 (10): 1683–99,
doi:10.3732/ajb.91.10.1683 ,
PMID 21652317 , retrieved
2011-01-28
35. Rothwell, G.W. & Nixon, K.C. (2006),
"How Does the Inclusion of Fossil
Data Change Our Conclusions about
the Phylogenetic History of
Euphyllophytes?", International
Journal of Plant Sciences, 167 (3):
737–749, doi:10.1086/503298
36. Stevens, P.F., Angiosperm Phylogeny
Website - Seed Plant Evolution
37. Hilton, Jason & Bateman, Richard M
(2006), "Pteridosperms are the
backbone of seed-plant phylogeny",
Journal of the Torrey Botanical
Society, 133 (1): 119–168,
doi:10.3159/1095-
5674(2006)133[119:PATBOS]2.0.CO
;2 , ProQuest 204065782
38. Lindley, J. (1830), Introduction to the
Natural System of Botany, London:
Longman, Rees, Orme, Brown, and
Green, OCLC 3803812 , p. xxxvi
39. Banks, H.P. (1975), "Reclassification
of Psilophyta", Taxon, 24 (4): 401–
 413, doi:10.2307/1219491 ,
JSTOR 1219491
40. Kenrick, P. & Crane, P.R. (1997), The
Origin and Early Diversification of
Land Plants: A Cladistic Study,
Washington, D.C.: Smithsonian
Institution Press, ISBN 978-1-56098-
730-7
41. Novíkov & Barabaš-Krasni (2015).
Modern plant systematics. Liga-
Pres. p. 685.
doi:10.13140/RG.2.1.4745.6164 .
ISBN 978-966-397-276-3.
42. Hao, Shougang & Xue, Jinzhuang
(2013), The early Devonian
Posongchong flora of Yunnan: a
 contribution to an understanding of
the evolution and early
diversification of vascular plants ,
Beijing: Science Press, p. 366,
ISBN 978-7-03-036616-0, retrieved
2019-10-25
43. Anderson, Anderson & Cleal (2007).
Brief history of the gymnosperms:
classification, biodiversity,
phytogeography and ecology.
Strelitzia. 20. SANBI. p. 280.
ISBN 978-1-919976-39-6.
44. Pelletier (2012). Empire biota:
taxonomy and evolution 2nd ed.
Lulu.com. p. 354. ISBN 978-
1329874008.
45. Lecointre, Guillaume; Guyader, Hervé
Le (2006). The Tree of Life: A
Phylogenetic Classification . Harvard
University Press. p. 175 .
ISBN 9780674021839.
"hemitracheophytes."
46. Nishiyama, Tomoaki; Wolf, Paul G.;
Kugita, Masanori; Sinclair, Robert B.;
Sugita, Mamoru; Sugiura, Chika;
Wakasugi, Tatsuya; Yamada, Kyoji;
Yoshinaga, Koichi (2004-10-01).
"Chloroplast Phylogeny Indicates
that Bryophytes Are Monophyletic" .
Molecular Biology and Evolution. 21
(10): 1813–1819.
doi:10.1093/molbev/msh203 .
ISSN 0737-4038 . PMID 15240838 .
47. Gitzendanner, Matthew A.; Soltis,
Pamela S.; Wong, Gane K.-S.; Ruhfel,
Brad R.; Soltis, Douglas E. (2018).
"Plastid phylogenomic analysis of
green plants: A billion years of
evolutionary history" . American
Journal of Botany. 105 (3): 291–301.
doi:10.1002/ajb2.1048 . ISSN 0002-
9122 . PMID 29603143 .
48. Phylogenomic Evidence for the
Monophyly of Bryophytes and the
Reductive Evolution of Stomata
49. Štorch, Petr; Žárský, Viktor; Bek, Jiří;
Kvaček, Jiří; Libertín, Milan (May 28,
2018). "Sporophytes of
polysporangiate land plants from the
 early Silurian period may have been
photosynthetically autonomous".
Nature Plants. 4 (5): 269–271.
doi:10.1038/s41477-018-0140-y .
PMID 29725100 . S2CID 19151297 .
50. Pryer, K.M.; Schuettpelz, E.; Wolf,
P.G.; Schneider, H.; Smith, A.R. &
Cranfill, R. (2004), "Phylogeny and
evolution of ferns (monilophytes)
with a focus on the early
leptosporangiate divergences" ,
American Journal of Botany, 91 (10):
1582–98,
doi:10.3732/ajb.91.10.1582 ,
PMID 21652310 , retrieved
2011-01-29, pp. 1582–3
51. Boyce, C.K. (2005), "The evolutionary
history of roots and leaves", in
Holbrook, N.M. & Zwieniecki, M.A.
(eds.), Vascular Transport in Plants,
Burlington: Academic Press,
pp. 479–499, doi:10.1016/B978-
012088457-5/50025-3 , ISBN 978-0-
12-088457-5
52. Sahney, S.; Benton, M.J. & Falcon-
Lang, H.J. (2010), "Rainforest
collapse triggered Pennsylvanian
tetrapod diversification in
Euramerica", Geology, 38 (12):
1079–1082,
Bibcode:2010Geo....38.1079S ,
doi:10.1130/G31182.1
53. Beerling, D.J. & Fleming, A.J. (2007),
"Zimmermann's telome theory of
megaphyll leaf evolution: a
molecular and cellular critique",
Current Opinion in Plant Biology, 10
(1): 4–12,
doi:10.1016/j.pbi.2006.11.006 ,
PMID 17141552
54. Tomescu, A. (2009), "Megaphylls,
microphylls and the evolution of leaf
development", Trends in Plant
Science, 14 (1): 5–12,
doi:10.1016/j.tplants.2008.10.008 ,
PMID 19070531
55. Smith, A.R.; Pryer, K.M.; Schuettpelz,
E.; Korall, P.; Schneider, H. & Wolf,
 P.G. (2006), "A classification for
extant ferns" (PDF), Taxon, 55 (3):
705–731, doi:10.2307/25065646 ,
JSTOR 25065646 , archived from the
original (PDF) on 2008-02-26,
retrieved 2011-01-28
56. Rutishauser, R. (1999), "Polymerous
Leaf Whorls in Vascular Plants:
Developmental Morphology and
Fuzziness of Organ Identities",
International Journal of Plant
Sciences, 160 (6): 81–103,
doi:10.1086/314221 ,
PMID 10572024
57. Chapman, Arthur D. (2009), Numbers
of Living Species in Australia and the
 World. Report for the Australian
Biological Resources Study ,
Canberra, Australia, retrieved
2011-03-11
58. Taylor, T.N.; Taylor, E.L. & Krings, M.
(2009), Paleobotany, The Biology and
Evolution of Fossil Plants (2nd ed.),
Amsterdam; Boston: Academic
Press, ISBN 978-0-12-373972-8, pp.
508ff.

Bibliography

Wikispecies has information related

to Embryophyta.

Raven, P.H.; Evert, R.F. & Eichhorn, S.E.

(2005), Biology of Plants (7th ed.), New
 York: W.H. Freeman, ISBN 978-0-7167-
1007-3
Stewart, W.N. & Rothwell, G.W. (1993),
Paleobotany and the Evolution of Plants
(2nd ed.), Cambridge: Cambridge
University Press, ISBN 978-0-521-
38294-6
Taylor, T.N.; Taylor, E.L. & Krings, M.
(2009), Paleobotany, The Biology and
Evolution of Fossil Plants (2nd ed.),
Amsterdam; Boston: Academic Press,
ISBN 978-0-12-373972-8

Retrieved from
"https://en.wikipedia.org/w/index.php?
title=Embryophyte&oldid=1009620269"
 Last edited 5 days ago by Tom.Reding

Content is available under CC BY-SA 3.0 unless

otherwise noted.