Prepared for B.Sc. Part II Botany Hons. by Prof.(Dr.

) Manorma Kumari

Lyginopteris oldhamia
Class Cycadopsida
Order Pteridospermales
Family Lyginopteridaceae
Genus Lyginopteris
Species L.oldhamia

Lyginopteris is well known genus in the family Lyginopteridaceae. It was

earlier known as Calymmatotheca hoeninghausii. Williamson, Scott, Brongniart,
Binney, Potonie and Oliver and Scott described this genus in detail. It was found
in abundance in the coal ball horizon of Lancashire and Yorkshire. Various parts
of the plants were discovered and named separately. They were later found to be
different organs of the same plants Lyginopteris due to presence of similar type
of capitate gland on them (except roots). Various parts are:
Leaves Sphenopteris hoeninghausii
Stems Lyginopteris oldhamia
Petioles Rachiopteris aspera
Roots Kaloxylon hookeri
Seeds Lagenostoma lomaxi

Morphological Features
The stem of Lyginopteris oldhamia was long aerial and radially
symmetrical varying in diameter from 2mm to 4cm. It was frequently branched
and bore adventitious roots that were probably prop roots that supported the weak
stems. The leaves were spirally arranged and up to 0.5 metre long. The leaves
were bipinnate to tripinnate. The pinnae had pinnules or leaflets and were present
at right angles to the rachis. The rachis and petiole were studded with capitate
glands.

Lyginopteris oldhamia: A. Showing external character; B. Frond bearing pollen sac; C.

Frond bearing seeds.
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Anatomy
Stem:
The transverse sections of the stem show nearly circular outline. Outer
layer is epidermis, next to it is many layered cortex. Outer cortex contains radially
broadened fibrous strands that forms a vertical network. It provides mechanical
strength to the weak stem. The inner cortex consists of parenchymatous cells. The
pericycle present just inner to the cortex, consist of short cells and a number of
thick walled or sclerotic cells with dark contents. Five mesarch primary vascular
bundles separated by parenchymatous areas are present. The phloem is towards
the outer side in each vascular bundle and consists of irregularly arranged small
cells. Next to it, is the cambium that is well preserved in some specimens. The
xylem consists of tracheids. The protoxylem tracheids have spiral thickenings.
Centripetal tracheids of the metaxylem have multiseriate bordered pits while the
centrifugal tracheids are scalariform. The pith is large and parenchymatous. Some
thick-walled cells, called sclerotic nests are scattered throughout the pith. Several
distinctly mesarch leaf traces are quite prominent. One leaf trace traverses
through the petiole and branches into two parts to supply the forking rachis.
The secondary structure of the stems exhibits the presence of periderm just
inner to the two' cortical layers. The primary phloem gets crushed and inner to
this the cylinder of the secondary vascular tissue is present and it surrounds the
primary xylem. Several medullary rays are in this region. The leaf traces interrupt
the cylinder of secondary vascular tissue. Many large and small cells are
alternately present in the secondary phloem. The cambium and then secondary
xylem are present inner to the secondary phloem region. The secondary xylem
consists of large tracheids with multiseriate bordered pits arranged on the radial
walls.

(A): Lyginopteris oldhamia A.T.S. (B): L.S. of Primary vascular bundle

of primary stem

Leaf
The pinnules are covered with a distinct layer of upper and lower
epidermises. The upper epidermis is cutinised and the cuticle is very resistant and
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well preserved. There are well defined palisade and spongy tissues that made up
the mesophyll tissue. The stomata are restricted to the underside.

Lyginopteris oldhamia T.S. stem showing secondary structure

Part of T.S. Pinna of Sphenopteris hoeninghausii

Root
The transverse section of root is roughly circular in outline. Cortex is
differentiated into outer cortex and inner cortex. Outer cortex made up of 2-3
layers of thin walled cells with scanty content. The inner cortex is 4-6 layered
whose cells were full of dense contents. These are considered to be muscilaginous
in nature. The endodermis and pericycle are clearly observed. Root possess many
primary xylem strands alternating with an equal number of phloem strands. The
xylem is exarch. The pith is scanty. The branches of root develop opposite the
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protoxylem points. The large roots show a secondary wood. It is traversed by

large rays that always lie opposite the protoxylem groups.

T.S. Root of Lyginopteris oldhamia

All the parts of the plant except the root are covered by numerous stalked
glandular outgrowths. These glandular outgrowths are flask-shaped and may be
up to 3mm high. They are not supplied by any vascular strand and are, therefore,
considered as mere emergences. Each outgrowth bears an apical globular head
that consists of a number of thin-walled secretory cells. It ranges in diameter from
0.12-0.4mm. Below the secretory head is a stalk made up of several layers of
parenchyma cells. These are capitate glands.

Reproductive Structure
Lyginopteris oldhamia was heterosporous, and its ovules and seeds were
enclosed within well protective cupules. Kidston (1905) discovered the
impressions of Crossotheca in connection with the leaves of Sphenopteris
hoeninghausii and thus it was concluded that Crossotheca is the
microsporangium of Lyginopteris because the leaves of Lyginopteris were
described under the name Sphenopteris hoeninghausii. The ovules and seeds of
Lyginopteris were described under the name Lagenostoma lomaxi. Arnold (1947)
stated that the microsporangiate structures of Lyginopteris belongs to Telangium.
Some other workers have also shown doubts on the validity of Crossotheca being
the microsporangium of Lyginopteris.
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Microsporangium
Crossotheca hoeninghausii has been described as the microsporangia of
Lyginopteris. Six to seven bilocular, pendant microsporangia were present on
more or less peltate fertile pinnules called microsporophylls. Each
microsporangium was about 3mm in length and its dehiscence was longitudinal.
Microspores were present in the cavities of microsporangia, sometimes even in
tetrads. Each microspore had a thick rough exine and attained a diameter of 50 to
70 micron. They have tri-radiate markings (trilete). The sporangia lack annulus
and resemble those of Cycas. There is no evidence of pollen tubes and so sperm
must have been motile.

Lyginopteris oldhamia, part of a frond bearing microsporangia

Ovule and Seed

Olive and Scott (1903) describe Lyginopteris oldhamia seeds as
Lagenostoma lomaxii. Seed was a small, barrel-shaped structure measuring 5 to
6mm in length and about 4mm in width. Each seed was surrounded by a cupule
present at the end of the slender rachis (Jongmans, 1930). Rachis was also
covered by capitate glands. The seeds and the cupule are known only in
petrified condition.
The ovule is orthotropous with a stout integument. The integument is not
lobed and forms a single complete sheath penetrated at the apex by a narrow
micropyle. The integument is fused with the nucellus except near the micropylar
end where it is free from the nucellus and forms a canopy. This canopy part is
divided into nine loculi with a vascular bundle extending into each.
The integument shows two distinct regions, the external sclerotesta which
is hard and stony and an inner fleshy endotesta. External to the integument, the
ovule is partially enclosed by a lobed cupule which is studded with numerous
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glandular outgrowths. The nucellus apex has a hollow pollen chamber or

Lagenostome.

V.S. of Ovule of Lyginopteris oldhamia

The seed of Lyginopteris oldhamia

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