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Origin, Characteristics and affinities of
Cycadofilicales:
The Cycadofilicales or Pteridosperms are popularly known as ‘seed
ferns.’ This group of Gymnosperms includes trees or small plants bearing
fern-like foliage. They flourished greatly in the Carboniferous
period(359mya----299mya), also known as the ‘Age of Ferns.’ The plants first
appeared in the Devonian, became very prominent in the Carboniferous, but
remained widespread during the Permian, but a few survived even up to
Mid-Jurassic. Grand Eury discovered them in 1877. The best-known
Cycadofilicales were present in Great Britain.
The presence of fern-like leaves was formerly believed to be a kind of fern,
but subsequently, after discovering fossil leaves with attached seeds in 1903,
Oliver and Scott designated the Cycadofilicales as Pteridosperms. They are
regarded as the primitive group of seed plants. Combined characters of both
ferns and cycads were observed, which led to the creation of a new
systematic group, the Cycadofilicales.
In the year 1877, Grand Eury first discovered a few species of fern leaves
which later on went by the name of the form genus Myeloxylon. In 1887
Williams pointed out that Heterangium (stem) and Kaloxylon (root)
combined characters of both ferns and cycads, which led Potonie (1899) to
create a new systematic group the Cycadofilicales.
With the discovery of the fact that Lagenostoma lomaxi (Lagenostoma is a
genus of seed ferns, based on ovules preserved in coal balls from the Six-Inch
Coal of the Hough Hill Colliery near Stalybridge, England) belongs to the stem
of Lynginopteris oldhamia by Oliver and Scott in 1903, the name
Pteridospermae was proposed for the seed-bearing Cycadofilicales. The
seed-bearing habit of the Pteridospermae was again verified next year by
Kidston in Neuropteris heterophylla.
Characteristic Features of Cycadofilicales:
1. Plants were trees or small plants with monopodial branching and the
presence of ramentum (rows or lines).
2. Plants possessed slender stems with large frond-like leaves, as in
Alethopteris & Sphenopteris.
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3. The primary xylem was mesarch, represented by solid or medullated
protostele. Polystelic
conditions were common in some members.
4. The secondary wood was manoxylic (loose and soft) and limited in
amount.
5. The cortex was well-developed and had a longitudinally aligned fiber
strand.
6. Leaves were usually fern-like (frond), spirally arranged, large, and
pinnately compound.
7. A resistant cuticle covered the leaves.
8. Young leaves are with circinate vernation.
9. Ovules are borne separately along margins or on the surface of
compound megasporophylls.
10. Megasporophylls did not arrange to form strobili.
11. Microsporophyll’s pinnately compound and not in strobili.
12. Microsporangia are without annulus and grouped into synangia.
13. A well-developed vascular supply was present in the seed.
14. The seeds were also provided with a definite pollen chamber.
15. Seeds are without annulus and are not in cones/inflorescence.
16. Seeds were on slightly modified or undistinguished foliage
17. The seeds resemble those of the present-day Cycads.
18. A cupule-like structure surrounds the ovules.
19. Pollen grains are monolete or trilete (Fig).
20. Pollination occurs without any pollen tube formation.
21. Male gametes are motile. They are directly discharged into the
pollen chamber.
Classification of Cycadofilicales:
The Cycadofilicales have been divided into two major groups:
The Paleozoic Cycadofilicales
1. Lyginopteridaceae
2. Medullosaceae
3. Calamopityaceae
The Mesozoic Cycadofilicales.
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1. Peltaspermaceae
2. Corystospermaceae
3. Caytoniaceae
Of the six families mentioned above, the most extensively studied one is
Lyginopteridaceae.
1. Lyginopteridaceae:
In the family, the leaves are of filicoid type (Fern-like), and the stem is of
monostelic organization. The leaf,
traces, and foliar bundles are
single or double; the xylem is
mesarch. Seeds are of cycadean
type, and the cupule is present.
The best-known lyginopterid is
Calymmatotheca hoeninghausi.
This is the plant to which both
Lyginopteris and Lagenostoma
belong.
2. Medullosaceae:
In this family, the stem is
polystelic, and the seeds are large.
3. Calamopityaceae:
It is the oldest and the largest of all the families of Pteridosperms. The stem
is monostelic, and scattered bundles are present in the petiole.
4. Peltaspermaceae:
The leaves in this family are small and bipinnate, and swellings characterize
the rachis. The microsporophyll is branched, microsporangia open
longitudinally, and smooth pollens are discharged.
5. Corystospermaceae:
The leaves in this family are linear-lanceolate. The seeds have oval, and
elliptic shapes and curved bifid micropyle.
6. Caytoniaceae:
This family is characterized by leaves formed usually in two (occasionally
three) pairs. Anatomically it has transfusion tissue and mesophyll with a
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palisade layer. The pollen grains show a symmetrical bilobed structure, and
wings.
The ovules are orthotropous, and the nucellus is free from the integument
except at the base. This is an advanced character. Formerly the Caytoniales
were compared with the Angiosperms, but later on, it was found out that
members of this family are essentially gymnospermous.
Affinities of Pteridospermales:
Pteridospermales (Cycadofilicales) have an intermediate position between
ferns and Cycadophytes due to their several resemblances with both groups.
But due to the presence of exposed seeds, they certainly belong to
gymnosperms.
Some of their possible affinities with ferns and Cycadophytes are;
Similarities with Ferns:
1. Large and pinnately compound leaves.
2. Young, circinately coiled leaves.
3. Mesarch or rarely exarch xylem.
4. Absence of vessels in the xylem and companion cells in the phloem.
5. The polystelic condition of Medullosa resembles ferns.
6. The sporangia were borne on the foliage leaves.
7. Motile flagellate Antherozoids.
8. Except for the presence of secondary wood, the vascular anatomy of the
stem is similar to that of ferns.
9. Presence of a thick wall around the megaspores.
Similarities with Cycadophytes (Cycadales):
Some common characteristics found in both Cycadales and Pteridospermales
are;
1. Both Cycadales and Pteridospermales possess pinnately compound
leaves with circinate vernation.
2. The wood is manoxylic in both groups.
3. Mucilage canals are present in the cortex and pith of the stem of both
cycads and “seed ferns.”
4. The extensive cortex is present in most Cycadales and some
Pteridospermales (e.g., Medullosa).
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5. Centripetal xylem is present in some Cycadales and the vegetative organs
of some “seed ferns” or Pteridospermales.
6. Male gametes or spermatozoids are multi-ciliate and motile in both
groups.
7. In both Pteridospermales and some Cycadales (e.g., Cycas), the
megasporophylls remain spirally and loosely arranged and do not form a
compact cone.
8. In Calymmatotheca and Cycas, the ovules remain attached to the
proximal parts of the leaves, and their distal or tip regions remain sterile.
9. The seeds of Lagenostoma lomaxi of Pteridospermales remain
surrounded by an outer hard stony layer and an inner fleshy layer. They
resemble the seeds of cycads to some extent.
10. Both groups are heterosporous.
11. The vascular supply of the ovules of cycads and some Pteridospermales
also show some similarities with each other.
12. A pollen chamber is present in the ovules of both these groups.
Similarities with Ginkgoes:
Ginkgo resembles the pteridosperms in having the following characteristics:
1. The leaf of Ginkgo looks like the wedge-shaped pinnule of pteridosperm.
2. There is an
accumulation of
abundant reserve food
material and
lignification in the integument before
fertilization.
3. The collar in the ovule of Ginkgo is similar to the
cupule of the pteridosperm ovule.
