Phylum: Anthocerotophyta

The hornworts that include 11 genera & more than 300 species constitute the
least diverged lineage of bryophytes. The name 'hornwort' is derived from their
horn like sporophytes.
1. The vegetative gametophyte of hornworts is a flattened thallus, with or
without a mid rib. Growing regions that contain solitary apical cells and
immediate derivatives typically are located in thallus notches and are
covered by mucilage that is secreted by epidermal cells.
2. The thickened thallus of the hornwort gametophyte lacks internal
differentiation except for the occurrence of rather extensive schizogenous
mucilage canals in species of Anthoceros.
3. Unlike in the sporophyte, all epidermal cells of the gametophyte contain
chloroplasts. Mucilage filled cells are abundant and scattered among
photosynthetic parenchyma in most taxa. Band like wall thickenings and
primary pit fields may occur in thallus cells subtending archegonia and
later the sporophyte foot. Cells contain solitary chloroplast with central
pyrenoids and channel thylakoids, features shared with algae but found
in no other land plants. Thylakoids traverse the pyrenoid and separate
typically lens-shaped subunits, giving the appearance of “multiple
pyrenoids”.
4. A distinctive feature of anthocerotes is the occurrence of apically derived
mucilage clefts on the ventral thallus through which Nostoc enters the
plant and becomes established as a colonial endosymbiont. Two cells
that resemble guard cells surround the opening which, once formed,
lacks the ability to open and close.
5. Rhizoids are unicellular, smooth and may have branched tips. They are
typically ventral in position and occasionally they develop from the outer
cell derived from a periclinical division of an epidermal cell.
6. Some hornworts have mycorrhiza like associations with glomerocytes.

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7. Asexual reproduction is widespread in anthocerotes. Fragmentation,
regenerant formation and gemmae production have been reported in
various taxa. Under adverse conditions, some species of hornworts
produce nutrient filled tubers as perennial bodies.
8. Most hornwort gametophytes are unisexual. In bisexual gametophytes,
development of the antheridia usually precedes that of archegonia.
Gametangia are produced along the dorsal thallus midline. Archegonia
are exogenous i.e. they develop from the surface cells, and ultimately
they are sunken in thallus tissue. In addition to the central cells of the
archegonium, the archegonial initial gives rise to one – or two layered
venter and six rows of neck cells that slightly protrude from the thallus
surface and overarched by a layer of mucilage. Venter cells are smaller
than the surrounding parenchyma.
9. Antheridia are referred endogenous because they develop from
subepidermal cells and ultimately are positioned within the internal
thallus chambers. In other embryophytes, antheridia develop from single
epidermal cells. The difference in hornworts is that the antheridial intial
is located at the base of a schizogenous antheridial chamber, and not at
the thallus surface. One to 25 antheridia are enclosed in sunken
chambers of the thallus.
10. Spermatids develop in pairs and the mature spermatozoids is coiled,
biflagellated and symmetrical. Both flagella insert at the anterior extreme
of the cell and are directed posteriorly. Spermatozoids are extremely
small (approximately 3.0μm in diameter) and only contain an anterior
mitochondrion, a cylindrical nucleus with mid constriction, and a
posterior mitochondrion associated with a plastid containing one starch
grain. Spermatozoids exhibit a right-handed (dextral) coil.
11. The first division of zygote is longitudinal and the endothecium of
embryo gives rise to a central columella in the sporophyte, if one exists.
This in contrast to most mosses and liverworts in which zygote
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 undergoes a transverse first division and the endothecium gives rise to
sporogenous tissue (exception Sphagnum). The foot matures before the
remaining histogenic regions. Growth of the overarching gametophytic
tissue occurs as the embryo develops, thus forming a protective involucre
that in most taxa is ruptured with continued maturation of sporophyte.
The involucre remains as cylinder that surrounds the base of the
sporophyte.
12. Multiple sporophytes are produced by each fertile thallus. At maturity
the sporophyte is differentiated into a bulbous foot embedded in
gametophyte tissue and a prominent elongated cylindrical spore-bearing
region which includes an epidermis, assimilative layer, sporogenous
tissue, columella and basal meristem. This meristem is located above the
foot and is responsible for the continuous production of sporogenous
tissue throughout the growing season.
13. Spore maturation occurs progressively from the base to the apex of the
sporophyte.
14. Stomates that resemble those of mosses and tracheophytes occur in the
sporophyte of many taxa. Guard cells are characterized by inner (ventral)
wall thickenings and apparently they are only epidermal cells contain
plastids, specifically amyloplastids. These features suggest homology
with stomates of other embryophytes. Epidermal cells are elongated and
may contain transverse annular wall thickening of unknown function.
15. At the foot placental transfer cells that contain elaborate wall labyrinth
are restricted to the gametophyte tissue, a feature that is shared with
Coleochaete and rare in other bryophytes. A distinctive feature of the
hornwort placenta is the occurrence of abundant protein crystals
between gametophyte and sporophyte cells in several genera.
16. An assimilative (photosynthetic) layer of variable thickness underlies
the epidermis and the sporogenous tissue is situated between this layer
and the columella. Pseudoelaters are interspersed among the spores and
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 are multicellular or unicellular with smooth or spiraled wall thickenings.
The central columella is persistent and usually consists of 16 cells in
cross section.
17. Dehiscence typically occurs along two longitudinal lines that originate
near the sporophyte tip. Pseudoelators and the columella facilitate spore
separation and assist in dispersal.
Classification of Anthocerotopsida Hyvonen & Pippo (1993)
Order: - Anthocerotales
Family: - Anthocerotaceae e.g., Anthoceros, Folioceros,
Phaeoceros
Family: - Dendrocerotaceae e.g., Dendroceros, Megaceros
Order: - Notothylales
Family: - Notothyladaceae e.g., Notothylas

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