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In these three types, Grammatopteris, Tubicaulis, and
Botryopteris, we have monostelic plants, for the most part of very
small size, with leaf-traces varying in shape from the oblong band-
form in Grammatopteris, and the oval form of Botryopteris antiqua, to
the ω type represented in its most pronounced form by B. forensis.
In several species the stem stele is endarch. Our knowledge of the
leaves is very meagre: in B. forensis they were repeatedly branched
and apparently bore small fleshy pinnules; the sporangia, though
differing from those of recent ferns, may be compared with the
spore-capsules of Osmundaceae as regards the structure of the
annulus. The abundance of hairs on the stems and leaves of some
species, the tracheal sheath in the sporangium described by
Oliver[1131] as Tracheotheca (= Botryopteris?), and the apparent
absence of a large well-developed lamina, may perhaps be regarded
as evidence of xerophilous conditions.
II. Zygoptereae.
Corda[1132] proposed the generic name Zygopteris for petrified
petioles from the Permian of Saxony, included by Cotta in his genus
Tubicaulis, which he named T. primarius. Corda’s genus has been
generally used for petioles of Palaeozoic ferns characterised by a
vascular strand having the form of an H in transverse section (fig.
308, D). Since the generic name was instituted, information has
been obtained in regard to the nature of the stems which bore some
of the petioles of the Zygopteris type; and for other species of
Zygopteris, the stems of which are still unknown, new generic names
have been proposed. P. Bertrand[1133] retains Zygopteris for one
species only, Z. primaria. Fig. 308, D, shows the character of the
petiolar vascular strand; the chief points are the comparatively long
cross-pieces (antennae of P. Bertrand) inclined at an angle of 45° to
the plane of symmetry of the petiole axis, and the groups of
protoxylem elements shown by the white patches in fig. D. In this as
in other members of the Zygoptereae the main rachis of the leaf
gives off four sets of branches in pairs alternately from the right and
left side of the primary vascular axis. This method of branching of the
stele in the primary rachis of several members of the
Coenopterideae shows that the fronds bore pinnae laterally
disposed, in some cases in one row and in others in two rows on
each side of the rachis. In a typical fern frond, as represented by
recent and most fossil species, branching of the rachis occurs in the
plane of the frond, that is in the plane represented by the horizontal
arm of xylem in Zygopteris primaria connecting the two antennae or
cross-pieces. In the Zygoptereae the branches from the petiole
vascular axis lie in a plane at right angles to that of the frond; they lie
in the transverse and not in the horizontal plane. The two strands
shown in fig. 308, B, 4, have been formed by the division of a single
strand, 3, in the transverse plane (i.e. in the plane of the paper). As
Tansley[1134] points out, a type of branching superficially similar to,
though not identical with this, is seen in some recent species of
Gleichenia and Lygodium. In this connexion it is worthy of note that a
fern figured by Unger from Thuringia as Sphenopteris petiolata
Goepp[1135] bears pinnae in two rows on the rachis which are
characterised by repeated branching and by a very narrow lamina or
by slender naked axes; the occurrence of this form of frond in rocks
containing Clepsydropsis antiqua (fig. 308, A) suggests a possible
connexion between the petrified rachis and the impressions of the
leaves.
Fig. 308.
A. Clepsydropsis antiqua.
B. Etapteris Scotti.
C. Diplolabis forensis.
D. Zygopteris primaria.
E–G. Stauropteris oldhamia.
The white patches in the xylem in figs. B–G mark the position of
protoxylem elements.
(A, after Unger; B–G, after P. Bertrand.)
Fig. 309.
A. Diplolabis forensis.
B. Botryopteris forensis.
C, D. Corynepteris coralloides.
E. Schizopteris (Etapteris) pinnata.
(A, B, after Renault; C, D, after Zeiller; E, after Renault and Zeiller.)
The vascular strand of the rachis of Zygopteris primaria (fig. 308,
D) is simpler than that of most of the Zygoptereae and exhibits a
close resemblance to the type of strand described by Renault as
Diplolabis (fig. 308, C).
Diplolabis.
Renault[1136] instituted this genus for two species from the Culm
beds and Coal-Measures of France based on the structure of the
petioles. The stems are unknown. The main rachis has a stele
similar to that of Zygopteris primaria, but distinguished by its greater
similarity, in transverse section, to an X rather than to the letter H:
the long transverse bar in Zygopteris is here much reduced in size.
The petiole of Diplolabis forensis[1137] Ren. (fig. 308, C) has a
diameter of 1·5–2 cm. From the antennae a pair of small bundles is
given off alternately from the right and left side, as in Zygopteris; the
members of each pair coalesce after leaving the antennae and then
separate to pass into the lateral branches of the frond. The position
of the protoxylem and the formation of the lateral xylem strands
previous to their separation are shown in fig. 308, C. On the side of
the vascular strand shown in fig. C, 2, the two lateral extensions of
the antennae are converging towards one another previous to their
separation and subsequent union. The ovoid sporangia occur in
groups of three to six and are coalescent below with a central
receptacle; they have no annulus, but the cells on the side next the
receptacle are smaller than those on the external wall (fig. 309, A).
The synangial form of the sorus suggests comparison with
Marattiaceae.
The species described by Renault from the Culm of Esnost is
regarded by P. Bertrand as identical with that described by Solms,
from the Culm of Falkenberg, as Zygopteris Roemeri[1138]. Diplolabis
is compared by P. Bertrand with Metaclepsydropsis, the generic
name given to the Lower Carboniferous petiole described by
Williamson as Rachiopteris duplex[1139].
Mr Gordon has recently described in a preliminary note a new type
of stem stele under the name Zygopteris pettycurensis from the
Lower Carboniferous plant bed of Pettycur[1140]: he regards the
petioles attached to the stem as identical with Zygopteris Roemeri
Solms-Laubach[1141]. This species, founded by Solms-Laubach on
petioles only, is placed by Bertrand[1142] in the genus Diplolabis and
regarded as identical with D. esnostensis Ren. The stele found by Mr
Gordon may therefore be assigned to the genus Diplolabis: it
includes two regions composed exclusively of tracheae and is
cylindrical in transverse section. The inner xylem zone consists of
short, square-ended, reticulately pitted elements and the outer zone
is composed of long and pointed conducting tracheae. The
scalariform protoxylem elements are situated between the two
metaxylem zones. As Mr Gordon says: this type of stem occupies a
position “in the Zygopteroid alliance” corresponding to that which
Thamnopteris Schlechtendalii (p. 329) occupies in the
Osmundaceous series. The discovery of this stem supplies another
link between the two fern groups, Osmundaceae and
Coenopterideae. Pelourde[1143] has described an imperfectly
preserved vascular strand from a locality near Autun as the type of a
new genus Flicheia esnostensis. Mr Gordon has pointed out to me
that this is a partially rotted petiole of Diplolabis esnostensis (=
Zygopteris Roemeri).
In their recent account of fossil Osmundaceous genera, Kidston
and Gwynne-Vaughan[1144] speak of the central parenchyma of the
existing medullated stele as being derived from tracheal tissue. They
add that if the Zygopteroid line of descent is at all close to the
Osmundaceous, we must be prepared for the existence of a
Zygopteris with a solid xylem like that of Thamnopteris: “such a
discovery, in fact, we hopefully anticipate[1145].” The new Pettycur
stem amply justifies this prophecy. It is noteworthy that Mr Gordon’s
stem affords an instance of the occurrence of a type of stele, similar
in its cylindrical form and in the absence of parenchyma to that of
Botryopteris, in a plant bearing leaves characterised by the
Zygopteris type of vascular strand.
Clepsydropsis.
Unger[1147] instituted this genus as a subdivision of Corda’s family
Rhaciopterideae[1148], the name having reference to the hour-glass
form of the vascular axis[1149]. The type-species C. antiqua (fig. 308,
A) is spoken of as the commonest fossil plant in the Devonian rocks
of Thuringia. In some sections the xylem has the form seen in fig.
308, A, in which an invagination of thin-walled tissue occurs at each
end; in other sections (fig. 308, A′) the bays become islands in the
xylem. Solms-Laubach speaks of Unger’s species as Rachiopteris
(Clepsydropsis) antiqua. P. Bertrand[1150], who has recently described
Unger’s plant, while recognising that C. antiqua and
Metaclepsydropsis duplex closely resemble one another, draws
attention to certain differences in the structure of the xylem which he
regards as sufficient to justify a generic separation. The leaf-traces
of Clepsydropsis are described by Bertrand as almost circular closed
rings of xylem instead of an arc as in Metaclepsydropsis.
Fig. 310.
A. Metaclepsydropsis duplex.
B, C. Stauropteris oldhamia.
D. Ankyropteris scandens.
[A, from a section in Dr Kidston’s Collection (Lower Carboniferous); B, C,
from sections in the Cambridge Botany School; D, after Stenzel.]
Ankyropteris.
Stenzel adopted this name for a subdivision of Corda’s genus
Zygopteris, applying it to a species described by Renault as Z.
Brongniarti, to a Permian species described by himself as Z.
(Ankyropteris) scandens, and to Z. Lacattii Ren.; Rachiopteris Grayi
Will. and Rachiopteris corrugata Will. are also included in this genus.
The characters emphasised by Stenzel[1151] are (i) the double anchor-
like form of the H-shaped petiole strand in which the lateral arms
(antennae) are curved like the flukes of an anchor, and (ii) the
emission of four rows of branches instead of two. The latter
distinguishing feature no longer holds good, as Z. primaria also gives
off four rows of bundles and not two as Stenzel supposed. P.
Bertrand has adopted Stenzel’s genus in a narrower sense[1152].
Fig. 312.
A. Thamnopteris Schlechtendalii. Leaf-trace: px, protoxylem; s,
island of parenchyma. (After Kidston and Gwynne-Vaughan.)
B. Ankyropteris corrugata. Single trachea with tyloses.
C. A. bibractensis. Part of foliar strand. (After P. Bertrand.)
Bertrand includes in Ankyropteris Renault’s species Zygopteris
bibractensis[1157] and Williamson’s species Rachiopteris
[1158]
corrugata : the former he names A. bibractensis var.
westphaliensis. The fossil described by Williamson as R. irregularis
or inaequalis[1159] are the secondary branches of A. bibractensis.