II. SPHENOPHYLLALES.
_Sphenophyllum._
The genus _Sphenophyllum_ is placed in a special class, as representing
a type which cannot be legitimately included in any of the existing
groups of Vascular Cryptogams. Although this Palaeozoic genus possesses
points of contact with various living plants, it is generally admitted
by palaeobotanists that it constitutes a somewhat isolated type among
the Pteridophytes of the Coal-Measures. Our knowledge of the anatomy
of both vegetative shoots and strobili is now fairly complete, and the
facts that we possess are in favour of excluding the genus from any of
the three main divisions of the Pteridophyta.
In Scheuchzer’s _Herbarium Diluvianum_ there is a careful drawing of
some fragments of slender twigs, from an English locality, bearing
verticils of cuneiform leaves, which the author compares with the
common _Galium_[860]. As regards superficial external resemblance, the
_Galium_ of our hedgerows agrees very closely with what must have been
the appearance of fresh green shoots of _Sphenophyllum_.
A twig of the same species of _Sphenophyllum_ is figured by
Schlotheim[861] in the first part of his work on fossil plants;
he regards it as probably a fragment of some species of Palm.
Sternberg[862] was the first to institute a generic name for this
genus of plants, and specimens were described by him in 1825 as a
species of the genus _Rotularia_. The name _Sphenophyllites_ was
proposed by Brongniart[863] in 1822 as a substitute for Schlotheim’s
genus, and in a later work[864] the French author instituted the genus
_Sphenophyllum_. Dawson[865] was the first to make any reference to
the anatomy of this genus; but it is from the examination of the much
more perfect material from St. Étienne, Autun, and other continental
localities, the North of England and Pettycur in Scotland, by Renault,
Williamson, Zeiller and Scott, that our more complete knowledge has
been acquired.
The affinity of _Sphenophyllum_ has always been a matter of
speculation; it has been compared with Dicotyledons, Palms, Conifers
(_Ginkgo_ and _Phyllocladus_), and various Pteridophytes, such as
_Ophioglossum_, _Tmesipteris_, _Marsilia_, _Salvinia_, _Equisetum_ and
the Lycopodiaceae[866].
[Sidenote: DEFINITION.]
We may define the genus _Sphenophyllum_ as follows:—
Stem comparatively slender (1·5–15 mm.?), articulated, usually somewhat
tumid at the nodes; the surface of the internodes is marked by more or
less distinct ribs and grooves which do not alternate at the nodes,
but follow a straight course from one internode to the next. A single
branch is occasionally given off from a node. Adventitious roots are
very rarely seen, their surface does not show the ridges and grooves of
the foliage-shoots.
The leaves are borne in verticils at the nodes, those in the same whorl
being usually of the same size, but in some forms two of the leaves are
distinctly smaller than the others. Each verticil contains normally
6, 9, 12, 18 or more leaves, which are separate to the base and not
fused into a sheath; the number of leaves in a verticil is not always
a multiple of six. They vary in form from cuneiform with a narrow
tapered base, and a lamina traversed by several forked veins, to narrow
uninerved leaves and leaves with a lamina dissected into dichotomously
branched linear segments. The leaves of successive whorls are
superposed.
The strobili are long and narrow in form, having a length in some
cases of 12 cm., and a diameter of 12 mm.; they occur as shortly
stalked lateral branches, or terminate long leaf-bearing shoots. The
axis of the cone bears whorls of numerous linear lanceolate bracts
fused basally into a coherent funnel-shaped disc, bearing on its upper
surface sporangiophores and sporangia.
The strobili are usually isosporous, but possibly heterosporous in some
forms.
The stem is monostelic, with a triarch or hexarch triangular strand
of centripetally developed primary xylem, consisting of reticulate,
scalariform and spiral tracheae; the protoxylem elements being
situated at the blunt corners of the xylem-strand. Foliar bundles are
given off, either singly or in pairs, from each angle of the central
primary strand. The secondary xylem consists of radially disposed
reticulate or scalariform tracheae, developed from a cambium-layer.
The phloem is made up of thin-walled elements, including sieve-tubes
and parenchyma. Both xylem and phloem include secondary medullary
rays of parenchymatous cells. The cortex consists in part of fairly
thick-walled elements; in older stems the greater part of the cortical
region is cut off by the development of deep-seated layers of periderm.
The roots are apparently diarch in structure, with a lacunar and smooth
cortex.
• • • • •
The branch of _Sphenophyllum emarginatum_ Brongn. given in fig. 109
shows the characteristic appearance of the genus as represented by this
well-known species which Brongniart figured in 1822. The Indian species
shown in fig. 111 illustrates the occurrence of unequal leaves in the
same whorl, and in fig. 110, _B_, we have a form of verticil in which
the leaves are deeply divided into filiform segments. A larger-leaved
form is represented by _S. Thoni_, Mahr. (fig. 110, _A_), a species
occasionally met with in Permian rocks.
No specimens of _Sphenophyllum_ have so far been found attached to
a thick stem; they always occur as slender shoots, which sometimes
reach a considerable length. One of the longest examples known is in
the collection of the Austrian Geological Survey; the axis is 4 mm. in
breadth and 85 cm. long, bearing a slender branch 61 cm. in length. The
manner of occurrence of the specimen as a curved slender stem on the
surface of the rock suggests a weak plant, which must have depended
for support on some external aid, either water or another plant. The
anatomical structure and other features do not favour the suggestion
of some writers that _Sphenophyllum_ was a water-plant[867], but there
would seem to be no serious obstacle in the way of regarding it as
possibly a slender plant which flung itself on the branches and stems
of stronger forest trees for support.
A. The anatomy of Sphenophyllum.
The following account of the structural features of the stem and root
is based on the work of Renault[868], Williamson[869] and Williamson
and Scott[870]. We may first consider such characters as have been
recognised in different examples of the genus, and then notice briefly
the distinguishing peculiarities of two well-marked specific types.
_a._ _Stems._
i. Primary structure.
In a transverse section of a young _Sphenophyllum_ stem such as that
diagrammatically sketched in fig. 105, _A_, we find in the centre
the xylem portion of a single stele with a characteristic triangular
form. The primary xylem consists mainly of fairly large tracheae with
numerous pits on their walls; towards the end of each arm the tracheids
become scalariform, and at the apex there is a group of narrower
spiral protoxylem elements. In the British species there is a single
protoxylem group at the apex of each arm, but Renault has described
some French stems in which the stele appears to be hexarch, having two
protoxylem groups at the end of each of the three rays of the stele.
The primary xylem strand of _Sphenophyllum_ has therefore a root-like
structure, the tracheids having been developed centripetally from the
three initial protoxylem groups. This type of structure is typical
of roots, but it also occurs in the stems of some recent Vascular
Cryptogams.
[Illustration: FIG. 104. Diagrammatic longitudinal section of
_Sphenophyllum_.
_c_, outer cortex; _b_, space next the stele, originally occupied
by phloem _etc._; _a_, xylem strand. (After Renault[871].) × 7.]
As a rule the tissue next the xylem has not been petrified, but in
exceptionally well-preserved examples it is seen to consist of a band
of thin-walled elements, of which those in contact with the xylem may
be spoken of as phloem, and those beyond as the pericycle. Succeeding
this band of delicate tissue there is a broader band of thicker-walled
and somewhat elongated elements, constituting the cortex. The specimen
drawn in fig. 105, _A_, shows very prominent grooves in the cortex
opposite the middle of each bay of the primary wood. It is these
grooves that give to the ordinary casts of _Sphenophyllum_ branches
the appearance of longitudinal lines traversing each internode. In a
longitudinal section of a stem, the cortical tissue (fig. 104, _c_)
is found to be broader in the nodal regions, thus giving rise to the
tumid nodes referred to in the diagnosis. The increased breadth at the
nodes does not mean that the xylem is broader in these regions, as it
is in Calamite stems. Small strands of vascular tissue are given off
from the three edges of the triangular stele (fig. 105 _A_) at each
node; these branch in passing through the cortex on their way to the
verticils of leaves. The space _b_ in the diagrammatic section of fig.
104 was originally occupied by the phloem and inner cortex. In some
species of _Sphenophyllum_ the apex of each arm of the xylem strand,
as seen in transverse section, is occupied by a longitudinal canal
surrounded by spiral tracheids, as in the primary xylem of the old stem
shown in fig. 105, _C_.
[Illustration: FIG. 105. _Sphenophyllum._
_A._ Transverse section of young stem.
_B._ Transverse section of the wood of a young stem; _px_,
protoxylem; _x_, secondary xylem. (_A_ and _B_. _Sphenophyllum
plurifoliatum._) × 20.
_C._ Transverse section of an old stem; (_S. insigne_); _a_,
phloem; _b_, periderm; _c_, fascicular secondary xylem;
_d_, interfascicular secondary xylem. × 9. (No. 914 in the
Williamson Collection.)
_D._ Longitudinal section of the reticulate tracheae and medullary
rays; _r_, _r_, _r_, of _S. plurifoliatum_. × 36.
_E._ Similar section of _S. insigne_. × 75. (_D_ and _E_ after
Williamson and Scott.)]
ii. Secondary structure.
With the exception of very young twigs the petrified _Sphenophyllum_
stems usually show a greater or less development of secondary wood.
In the xylem-strand of fig. 105, _B_, the broad concave bays of the
primary wood have been filled in by the development of two rows of
large secondary tracheids, _x_, but opposite the protoxylem groups,
_px_, there are no signs of cambial activity. In the unusually large
stem represented by a rough sketch in fig. 105, _C_, the triangular
primary xylem lies in the centre of a thick mass of secondary vascular
tissue. The secondary and primary wood together have a diameter of
about 5 mm.
After the bays between each protoxylem corner have been filled
in, the formation of secondary wood proceeds uniformly along the
stem radii, but the rows of tracheids and medullary rays which are
developed opposite the corners of the primary strand, _c_, differ in
certain characters from the broader masses of wood opposite the bays.
For convenience, the secondary wood, _c_, opposite the protoxylem
groups has been spoken of as _fascicular wood_, and the rest, _d_, as
_interfascicular wood_.
The secondary xylem consists either of tracheae with numerous bordered
pits on their radial walls (fig. 105, _D_), or of tracheae with broad
and bordered scalariform pits (fig. 105, _E_). The suggestion of
concentric rings of growth in the wood in fig. 105, _C_, is rather
deceptive; there are no well-marked regular rings in _Sphenophyllum_
stems, but irregular bands of smaller elements occasionally interrupt
the uniformity of the secondary xylem. In some stems the medullary rays
have the form of rows of parenchymatous cells, which in tangential
longitudinal section are found to consist frequently of a single row
of radially disposed elements; this type of medullary rays occurs
in the species _Sphenophyllum insigne_, in which the tracheae are
scalariform. Three medullary rays, _r_, are seen on the radial face of
the scalariform tracheids in fig. 105, _E_, which represents a radial
section of this species. In other species, _e.g._ _S. plurifoliatum_,
the medullary rays have a peculiar and characteristic structure;
in a transverse section of the stem they appear as groups of a few
parenchymatous cells in the spaces between the truncated angles of the
large tracheae (fig. 106). In longitudinal section these medullary-ray
elements resemble thick bars stretching radially across the face of the
tracheae (fig. 105, _D_, _r_); the apparent septa or bars are however
thin-walled cells connecting the different groups of medullary-ray
cells, as seen in a transverse section. These radial connecting cells
are occasionally seen as short rays in transverse sections of stems.
The cambium and phloem elements are occasionally preserved in good
specimens of older stems; the former consist of tabular flatted
thin-walled cells, and the latter in some cases include large
sieve-tubes and narrower parenchymatous elements.
The sections shown in fig. 107, _E_ and _F_, illustrate the
preservation of cambial and phloem tissue. In the transverse section
of fig. 107, _F_, the secondary xylem with the medullary rays, _r_, is
succeeded by a few tabular cambium cells, and external to these there
are thin-walled elements of unequal size representing the phloem.
In fig. 107, _E_, the scalariform tracheids are succeeded by narrow
thin-walled cells, and the larger elements with transverse and oblique
septa are no doubt sieve-tubes.
In the large stem of fig. 105, _C_, the xylem is succeeded by
a band of tissue, a, which is no doubt phloem, and external to
this there is a considerable development of periderm (_b_). The
periderm in _Sphenophyllum_ stems had a deep-seated origin, the
phellogen or cork-cambium occasionally being formed in the secondary
phloem-parenchyma, and in other cases in the pericycle, as in the stems
of some living dicotyledons. Williamson and Scott[872] describe stems
in which a succession of phellogens were formed at different levels,
thus producing a scaly type of bark, such as we find in the Pine or the
Plane tree.
[Sidenote: SPHENOPHYLLUM PLURIFOLIATUM.]
Before describing the structure of the strobili of _Sphenophyllum_, we
may briefly point out the distinguishing features of two specific types
of the genus recently described by Williamson and Scott. One of these
species, _S. insigne_, was originally described by Williamson as an
_Asterophyllites_; the numerous narrow linear leaves in each verticil
led to the inclusion of the specimens in the latter genus. The material
on which this species is founded is from the volcanic beds of Pettycur,
Burntisland, on the coast of the Firth of Forth.
1. _Sphenophyllum insigne_ (Will.). Figs. 105, _C_ and _E_,
and 107, _E_ and _F_.
1891. _Asterophyllites insignis_. Williamson[873].
An _intercellular space_ occurs at each angle of the three-rayed
primary xylem strand, and spiral tracheae are abundant. The tracheae
of the secondary wood have _scalariform markings_ on the radial walls.
_Regular medullary rays_ extend through the secondary wood. The phloem
contains large sieve-tubes.
This species occurs in the Calciferous sandstone rocks of Burntisland,
and has lately been recorded from Germany. It characterises a lower
horizon than _S. plurifoliatum_ (Will. and Scott).
2. _Sphenophyllum plurifoliatum_ (Williamson and Scott)[874].
Figs. 105, _A_, _B_, and _D_, and 106.
1891. _Asterophyllites sphenophylloides._ Will.[875]
The specific name _plurifoliatum_ was proposed by Williamson and
Scott for a type of stem originally described by Williamson[876] as
an _Asterophyllites_, from the Coal-Measures of Oldham, Lancashire.
This form of stem has not so far been connected with any of the
older species founded on external characters, but it evidently bore
foliage in which the leaves were deeply divided, as in _Sphenophyllum
trichomatosum_ (fig. 110, _B_).
[Illustration: FIG. 106. Sphenophyllum plurifoliatum, Will. and Scott.
From a photograph by Mr Highly from a section in the Williamson
Collection (no. 899). × 27.]
In this species there are _no canals_ at the angles of the primary
xylem, and there are fewer spiral tracheae than in _S. insigne_. The
tracheae of the secondary wood have _numerous small pits_ on the radial
walls, and the medullary rays are chiefly composed of parenchymatous
cells, which appear in transverse section as _groups of cells_ between
the truncated angles of the tracheae. The characters are fairly well
seen in the xylem portion of a stele shown in fig. 106. The fascicular
wood includes some rows of parenchymatous medullary-ray cells in
addition to the characteristic groups, as seen in the figure. A
slightly oblique transverse section of a stem is often convenient in
the interpretation of histological features; one of the sections of _S.
plurifoliatum_ in the Williamson collection (no. 893), which has been
cut somewhat obliquely, shows very clearly the differences in pitting
exhibited by the different xylem elements.
_b. Roots._
Our knowledge of the anatomy of _Sphenophyllum_ roots is very limited.
Renault has described a somewhat imperfect example of a silicified
root from St. Étienne and Autun. The drawing in fig. 107, _B_, which
is copied from one of Renault’s figures, shows a cylindrical mass of
xylem with a small band of narrower elements occupying the centre, and
surrounded by rows of larger secondary tracheae. The central bipolar
band is described as the diarch primary xylem, around which the
secondary pitted elements have been developed.
It is probable that the specimen described by Renault is a root of
_Sphenophyllum_, but my impression gained from an examination of the
section was that the diarch primary strand is not quite so clear as
in the published figures. Until we possess better material we cannot
attempt any very satisfactory description of the anatomical features of
the roots of this genus.
A section of a _Sphenophyllum_ stem has been figured by Felix[877], in
which a lateral member is being given off; this may possibly represent
the origin of an adventitious root, but the preservation is not
sufficiently distinct to render this certain.
_c. Leaves._
Renault[878] has described some silicified leaves of _Sphenophyllum_
from Autun in which the laminae consist of thin-walled loose
parenchyma, traversed by small groups of tracheids constituting the
simple or forked veins. The epidermis is made up of a single layer of
cells, with here and there indistinct indications of stomata. A more
perfect stoma has, however, been described by Solms-Laubach from the
epidermis of a bract in a strobilus (fig. 107, _A_).
[Illustration: FIG. 107. A. Stoma in a bract of _Sphenophyllostachys_.
B. Root of _Sphenophyllum_. C. _Sphenophyllostachys Römeri_, Solms.
_s_, sporangiophore, _b_, bract. D. Sporangium. E and F. Sections
through the cambium, phloem and secondary xylem of _Sphenophyllum
insigne_ (Will.). _s_, sieve-tube. G. Sporangium and pedicel. A, C,
D. After Solms-Laubach. B. After Renault. E–G. After Williamson and
Scott. E. F. × 100. G. × 115.]
_d. Cones._
The history of the recognition of the cones of _Sphenophyllum_ has
already been briefly alluded to in chapter V., p. 100. The main
points in the structure of the cones of this genus were known for
several years, before the fact was established that they belonged to
_Sphenophyllum_ stems. In 1871 Williamson[879] published an account
of an imperfect fossil strobilus from the Lower Coal-Measures of
Oldham, Lancashire, under the name of _Volkmannia Dawsoni_. The
generic term _Volkmannia_ has been used by different writers for
cones varying considerably in structural features; in the case of
Williamson’s fossil, Weiss[880] substituted the name _Bowmanites_,
a genus instituted by Binney[881] for a strobilus apparently of the
same type as _Volkmannia Dawsoni_. In 1891 Williamson[882] described
some additional specimens of _Bowmanites Dawsoni_, and, as in his
earlier paper, he compared the strobilus with _Asterophyllites_ and
_Sphenophyllum_, but it was still a matter of speculation as to what
was the form of the vegetative branches. Soon after the more complete
account of the English cones was published, Zeiller[883] recognised
a close agreement between some French and Belgian specimens of
_Sphenophyllum_ strobili and the strobilus described by Williamson.
A closer comparison thoroughly established the connection between
_Bowmanites Dawsoni_ and _Sphenophyllum_; and there is little doubt
that this strobilus belongs to the stem known as _Sphenophyllum
cuneifolium_ (Sternb.)—a well-known species of the genus.
[Sidenote: STROBILUS.]
The most important morphological features of the strobilus of
_Sphenophyllum_ may best be illustrated by a detailed account of one
specific type, and by a brief reference to other forms which are
characterised by certain differences in the number and attachment of
the sporangia. When we know that a given strobilus must have grown on
a _Sphenophyllum_ stem, the obvious name to assign to it would seem
to be that of the plant which bore it; but there are advantages in
making use of special generic terms for detached cones, which cannot
be referred with certainty to a particular species of stem. The genus
_Calamostachys_ affords an example of a name which is intended to
denote that a cone so called belongs to a Calamarian plant; similarly
such a name as _Sphenophyllostachys_ may be used for Sphenophylloid
cones which cannot be connected with certainty to particular species
of _Sphenophyllum_. It has been suggested that the genus _Bowmanites_,
first used for a cone which was afterwards recognised as belonging to a
_Sphenophyllum_, should be employed instead of the sesquipedalian term
_Sphenophyllostachys_. The latter is used here as being in accordance
with a generally accepted and convenient system of nomenclature, and
as a name which at once denotes the fact that the fossil is not only a
cone but that it belongs to a _Sphenophyllum_.
_Sphenophyllostachys Dawsoni_ (Will.). Figs. 107, _A_ and _G_, 108.
Probably the strobilus of _Sphenophyllum cuneifolium_ (Sternb.).
[Illustration: FIG. 108. Diagrammatic longitudinal section of a
_Sphenophyllum_ strobilus. The upper figure represents a portion of
a whorl of bracts. (The smaller figure, after Zeiller.)]
The cone consists of a central axis bearing a number of verticils
of bracts coherent in their lower portions in the form of a widely
open funnel-shaped disc, which splits up peripherally into 14–20
linear-lanceolate segments. The free segments of each verticil have an
obliquely ascending or almost vertical position, and extend upwards
for a distance of about six internodes. The smaller drawing in fig.
108 shows the appearance in side view of the narrow bracts of a single
whorl. A transverse section of a strobilus would include, therefore,
sections of several concentric series of ascending bracts. The
verticils of _Sphenophyllostachys Dawsoni_ are probably superposed,
but this point has not been definitely settled. From the upper surface
of the coherent basal portion of each verticil, there are given off
twice as many sporangiophores as there are free segments, and these
are attached close to the line of junction of the axis of the cone
and the funnel-shaped disc. Each sporangiophore has the form of a
slender stalk which bends inwards at its distal end and bears a single
sporangium (_cf._ fig. 107, _D_). The sporangiophores given off from
the same verticil of bracts vary in length. All the sporangiophores
are attached to the coherent bracts at the same distance from the axis
of the cone; but as the sporangia between each verticil of bracts
are arranged in two or three concentric series, it follows that the
length of the sporangiophores varies considerably. The diagrammatic
longitudinal section of a strobilus in fig. 108 shows three concentric
series of sporangia between successive bract-verticils. A similar
diagram was published by Williamson in 1892[884], and afterwards copied
by Potonié[885], but in Williamson’s restoration the sporangiophores
of the three series of sporangia are erroneously represented as
arising from different points on the surface of the bracts. There is
little doubt, as regards the strobilus of _S. cuneifolium_, that the
sporangiophores were given off in a single series close to the axils of
the bracts, as is partially shown in fig. 108.
The central part of the axis of the cone is occupied by a single
triangular stele like that of the stem, except that each ray of the
xylem strand has a comparatively broad blunt termination, and is
not tapered to a narrow arm as in fig. 105, _A_ and _B_. The wood
consists of pitted tracheae, with two groups of protoxylem elements at
each of the truncated angles of the solid strand of xylem. From the
angles of the stele branches of vascular tissue pass out through the
cortex to supply the sterile and fertile segments of each verticil.
One of the transverse sections of the _Sphenophyllum_ cone in the
British Museum Collection (no. 1898 _E_) affords a good example of the
misleading appearance occasionally presented by an intruded ‘rootlet’
of _Stigmaria_; the vascular tissue of the cone has disappeared, and a
Stigmarian appendage with its vascular bundle occupies the position of
the stelar tissues.
The bracts consist of parenchymatous tissue limited externally by an
epidermis containing stomata. A single stoma with subsidiary cells
is represented in fig. 107, _A_. The sporangiophores are composed
internally of thin-walled cells with stronger cells towards the
surface. The longer sporangiophores in a series may be more or less
coherent for part of their length to the upper surface of the verticil
of bracts. In fig. 108 the slender sporangiophores do not appear to
come off always from the same portion of the bracts, but this is due to
some of them lying on the surface of the latter during part of their
course to support the external circle of sporangia. The hook-like
distal end of a sporangiophore, towards the point of attachment of the
sporangium, is characterised by the larger size and greater prominence
of the surface cells; these larger cells, which pass over the upper
surface of a sporangium base, probably constitute a kind of _annulus_
which determines the dehiscence of the sporangial wall[886].
Fig. 107, _G_, represents a sporangiophore and its sporangium cut
through transversely just below the point of attachment of the latter
to the end of the hook-like termination of the former. The spores are
characterised by an irregularly reticulate thickening of the outer coat
or exospore, as seen in the figure.
One of the chief points of interest suggested by a _Sphenophyllum_
cone is the exact morphological nature of the sporangiophores. Are
they branches borne in the axils of bracts, or may we regard each
sporangiophore as a modified leaf, which has become coherent with
the whorls of sterile leaves? Or is a sporangiophore merely a stalk
of a sporangium; or a ventral lobe of a leaf, of which the sterile
bracts represent the dorsal lobes? Although it is impossible without
the evidence of development to decide with certainty between these
alternatives, it would seem most probable that a sporangiophore may
be looked upon as a ventral lobe of a leaf, the sterile lobes forming
the bracts or members of the sterile whorls of the cone. This question
is discussed by Zeiller[887] and Williamson and Scott[888], also more
recently by Scott[889] in his memoir on _Cheirostrobus_.
_Sphenophyllostachys Römeri_ (Solms-Laubach)[890].
Fig. 107, _C_ and _D_.
In another type of _Sphenophyllum_ strobilus, recently described
by Solms-Laubach, the incurved end of each sporangiophore bore two
sporangia. In most respects this species, which has not been found in
connection with a vegetative shoot, agrees with _Sphenophyllostachys
Dawsoni_.
In fig. 107, _C_, which is copied from one of Solms-Laubach’s
drawings[891], we have an oblique transverse section of part of a
strobilus, including portions of two series of sporangia borne on one
verticil of bracts, and at the right-hand edge the section has passed
through the sporangia belonging to another whorl of bracts. There
were probably three concentric series of sporangia attached to each
verticil of bracts, as in the case of fig. 108. The unshaded area,
_b_ (fig. 107, _C_), represents the bracts of two successive sterile
whorls in transverse section. The shaded areas are the sporangia, with
their sporangiophores, _s_. The relative position of the sporangia and
sporangiophores suggests that each pedicel bore two sporangia at its
tip, instead of one, as in the strobilus of _Sphenophyllum cuneifolium_
(Sternb.).
A further variation in the structure of the strobili is illustrated by
some specimens of _S. trichomatosum_ Stur, described by Kidston[892],
from the Coal-Measures of Barnsley. Each whorl of bracts bears a single
series of oval sporangia which appear to be sessile on the basal
portion of the whorl. It is possible that delicate sporangiophores
may have been present, but in the imperfect examples in Kidston’s
collection[893] the sporangia present the appearance of being seated
directly on the surface of the bracts. As the specimens do not show any
internal structure, it would be unwise to lay too much stress on the
apparent absence of the characteristic sporangiophores. In any case,
Kidston’s cones afford an illustration of the occurrence of a single
series of sporangia in each whorl, instead of the pluriseriate manner
of occurrence in some other species.
The statement is occasionally met with that some _Sphenophyllum_ cones
possessed two kinds of spores, but we are still in want of satisfactory
evidence that this was really the case. Renault has described an
imperfect specimen, which he considers points to the heterosporous
nature of a _Sphenophyllum_ cone, but Zeiller and Williamson and Scott
have expressed doubts as to the correctness of Renault’s conclusions.
While admitting the possibility of undoubted heterosporous strobili
being discovered, we are not in a position to refer to _Sphenophyllum_
as having borne strobili containing two kinds of spores[894].
[Sidenote: SPHENOPHYLLUM EMARGINATUM.]
[The following are some of the specimens in the Williamson Cabinet
which illustrate the structure of _Sphenophyllum_:—
_S. plurifoliatum._ 874, 882, 884, 893, 894, 897, 899, 901, 903,
908, 1893.
_S. insigne._ 910, 914, 919, 921, 922, 924, 926, 1420, 1898.
_Sphenophyllostachys._ 1049A–1049C, 1898.]
B. Types of vegetative branches of Sphenophyllum.
1. _Sphenophyllum emarginatum_ (Brongniart). Fig. 109.
1822. _Sphenophyllites emarginatus_, Brongniart[895].
1828. _Sphenophyllum emarginatum_, Brongniart[896].
1828. _Sphenophyllum truncatum_, Brongniart[896].
1828. _Rotularia marsileaefolia_, Bischoff[897].
1862. _Sphenophyllum osnabrugense_, Römer[898].
[Illustration: FIG. 109. _Sphenophyllum emarginatum_ (Brongniart).
From a specimen in the Collection of Mr R. Kidston, Upper
Coal-Measures, Radstock. ⅚ nat. size.]
This species of _Sphenophyllum_ bears verticils of six or eight
wedge-shaped leaves varying in breadth and in the extent of dissection
of the laminae; they are truncated distally, and terminate in a margin
characterised by blunt or obtusely-rounded teeth, each of which
receives a single vein. The larger leaves are usually more or less
deeply divided by a median slit. The narrow base of each leaf receives
a single vein which branches repeatedly in a dichotomous manner in
the substance of the lamina. Several drawings have been given by
Sterzel[899] in a memoir on Permian plants, showing the variation in
leaf-form in _Sphenophyllum emarginatum_, but as Kidston[900] and
Zeiller[901] have pointed out Sterzel’s specimens probably belong to
_S. cuneifolium_ (Sternb.).
Branches are given off singly from the nodes, and the cones are borne
at the tips of branches or branchlets. The cone of _S. emarginatum_
agrees very closely with that of _S. cuneifolium_, and is of the same
type as that shown in fig. 108. The small branch of _S. emarginatum_
represented in fig. 109 does not show clearly the detailed characters
of the species, as the leaf-margins are not well preserved.
In one of the largest specimens of this species which I have seen, in
the Leipzig Museum, the main stem has internodes of about 3·9 cm. in
length, from which a lateral branch with much shorter internodes is
given off from a node.
It is important to notice the close resemblance, as pointed out by
Zeiller, between some of the narrower-leaved forms of _S. emarginatum_
and _S. cuneifolium_ (Sternb.)[902]; but in the latter species the
margins of the leaves have sharp, and not blunt teeth.
The cone described and figured by Weiss[903] as _Bowmanites
germanicus_, since investigated by Solms-Laubach[904], must be referred
to this species. Geinitz[905] figured a cone in 1855 as that of
_S. emarginatum_, but his determination of the species is a little
doubtful. Good figures of the true cone of _S. emarginatum_ have been
given by Zeiller[906] in his _Flore de Valenciennes_, as well as in his
important memoir on the fructification of _Sphenophyllum_.
[Sidenote: LEAVES.]
2. _Sphenophyllum trichomatosum_ Stur. Fig. 110, _B_.
The finely-divided leaves of the single whorl shown in fig. 110, _B_
(from the Middle Coal-Measures of Barnsley, Yorkshire), afford an
example of a form of _Sphenophyllum_ which is represented by such
species as _S. tenerrimum_ Ett.[907], _S. trichomatosum_ Stur[908],
and _S. myriophyllum_[909] Crép. Probably the specimen should be
referred to _S. trichomatosum_, but it is almost impossible to speak
with certainty as to the specific value of an isolated leaf-whorl of
this form. It has long been known that the leaves of _Sphenophyllum_
may vary considerably, as regards the size of the segments, on the
same plant; and the occurrence of such finely-divided leaves has
lent support to an opinion which was formerly held by some writers,
that _Asterophyllites_ and _Sphenophyllum_ could not be regarded as
well-defined separate genera. This heterophylly of _Sphenophyllum_
has thus been responsible for certain mistaken opinions both as to
the relation of the genus to _Calamocladus_[910] (_Asterophyllites_),
and as regards the view that the finely-divided laminae belonged
to submerged leaf-whorls, while the broader segments were those of
floating or subaerial whorls.
There is a very close resemblance between some of the deeply-cut and
linear segments of a _Sphenophyllum_ and the leaves of _Calamocladus_,
but in the former genus the linear segments are found to be connected
basally into a narrow common sheath. The assertion[911] that the
deeply-cut leaves occur on the lower portions of stems is not supported
by the facts. Kidston[912] has pointed out that the cones are often
borne on branches with such leaves, and the same author refers to a
figure by Germar, in which entire and much-divided leaves occur mixed
together in the same individual specimen. M. Zeiller recently pointed
out to me a similar irregular association of broader and narrower
leaf-segments on the same shoots in some large specimens in the École
des Mines, Paris. Cones of _Sphenophyllum tenerrimum_ have been figured
by Stur[913] and others; they are characterised by their small size
and by the dissection of the slender free portions of the narrow
bracts[914].
3. _Sphenophyllum Thoni_ Mahr. Fig. 110, _A_.
Another type of _Sphenophyllum_ is illustrated by _S. Thoni_ Mahr as
shown in fig. 110, _A_. This species was first described by Mahr[915]
from the Coal-Measures of Ilmenau, and has since been figured by
Zeiller and other authors. Each whorl consists of six large obcuneiform
leaves with the broad margin somewhat irregularly fringed. The
unusually good specimen of which fig. 110, _A_, represents a single
verticil was originally described and figured by Zeiller in 1880[916];
it is now in the École des Mines Museum, Paris.
[Illustration: FIG. 110.
_A._ _Sphenophyllum Thoni_, Mahr. (After Zeiller.)
_B._ _Sphenophyllum trichomatosum_, Stur. From a specimen in the
Woodwardian Museum; from the Coal-Measures of Barnsley, Yorks.
_A_ and _B_ ¾ nat. size.]
The leaf-forms illustrated by figs. 109 and 110 are some of the more
extreme types of _Sphenophyllum_ leaves; but these are more or less
connected by a series of intermediate forms. For a more complete
systematic account of the different species the student should consult
such works as those by Coemans and Kickx[917], Zeiller, Schimper, and
others.
4. _Sphenophyllum speciosum_ (Royle). Fig. 111.
1834. _Trizygia speciosa_, Royle[918].
The species shown in fig. 111 has been usually described as a separate
genus _Trizygia_, a name instituted by Royle in 1834 for some Indian
fossils from the Lower Gondwana rocks of India[919]. Zeiller[920] has
lately pointed out the advisability of including this Asiatic type in
the genus _Sphenophyllum_. The slender stem bears verticils of cuneate
leaves in three pairs at each node, the anterior pair being smaller
than the two lateral pairs. The characteristic _Sphenophyllum_ venation
is clearly seen in the enlarged leaf, fig. 111, _B_.
[Illustration: FIG. 111. _Sphenophyllum speciosum_ (Royle).
_A._ Nat. size. _B._ enlarged leaf.
From the Raniganj Coal-field, India. (After Feistmantel.)]
The inequality of the members of a single whorl, which characterises
this Indian plant, is sometimes met with in European species. A
specimen of _Sphenophyllum oblongifolium_, which Prof. Zeiller showed
me in illustration of this point, was practically indistinguishable
from _Trizygia_[921].
In some of the earlier descriptions of the Indian species the generic
name _Sphenophyllum_[922] was used by McClelland and others, but the
supposed difference in the leaf-whorls was made the ground of reverting
to the distinct generic term _Trizygia_. Now that a similar type of
leaf-whorl is known to occur in _Sphenophyllum_, it is better to adopt
that genus rather than to allow the question of locality to unduly
influence the choice of a separate generic name for an Indian plant.
[Sidenote: GEOLOGICAL RANGE.]
C. Affinities, range and habit of Sphenophyllum.
It has been pointed out in the description of _Sphenophyllum_, that
the most widely separated families of recent plants have been selected
by different authors as the nearest living allies of this Palaeozoic
genus. It is now generally admitted that _Sphenophyllum_ is a generic
type apart; it cannot be classed in any family or sub-class of recent
or fossil plants, without considerably extending or modifying the
recognised characteristics of existing divisions of the plant-kingdom.
The anatomical characters of the _Sphenophyllum_ stem are such as one
finds in some recent genera of the Lycopodineae, especially _Psilotum_.
If the stele of _Psilotum_ were composed internally of a solid
strand of xylem, we should have a close correspondence between the
centripetally-developed wood of this genus and that of _Sphenophyllum_.
Similar comparisons might be drawn with other existing genera, but the
more detailed consideration of the affinities of the Palaeozoic plant
will be more easily dealt with after other members of the Pteridophytes
have been described. The recent discovery of an entirely new type of
Carboniferous strobilus in rocks of Calciferous sandstone age on the
shores of the Firth of Forth has thrown new light on the position of
_Sphenophyllum_. _Cheirostrobus Pettycurensis_, the new cone which
Scott has described in an able memoir, affords certain points of
contact with _Sphenophyllum_ on the one hand and with _Calamites_ on
the other. This important question will be dealt with after we have
given an account of _Cheirostrobus_[923]. To put the matter shortly,
_Sphenophyllum_ agrees with some Lycopodinous plants in its anatomical
features; with the Equisetales it is connected by the verticillate
disposition of the leaves, and some of the forms of _Sphenophyllum_
strobili present features which also point to Equisetinous affinities.
In his Presidential address to the Botanical Section at the
British Association Meeting of 1896 Scott[924] thus refers to the
Sphenophyllums:—“We may hazard the guess that this interesting group
may have been derived from some unknown form lying at the root of both
Calamites and Lycopods. The existence of the Sphenophyllae certainly
suggests the probability of a common origin for these two series.” The
result of the subsequent investigation of the new cone _Cheirostrobus_
amply justifies this opinion as to the position of _Sphenophyllum_.
It is probable that _Sphenophyllum_ lived during the Devonian period,
but the unsatisfactory specimens on which Dawson has founded a species
of this age, _S. antiquum_[925], can hardly be said to afford positive
evidence of the Pre-Carboniferous existence of the genus. From the
Culm rocks and other strata older than the Coal-Measures, we have
such species as _S. insigne_ (Will.), _Sphenophyllostachys Römeri_
(Solms-Laubach), and _Sphenophyllum tenerrimum_, Ett.[926] while _S.
emarginatum_[5], Brongn. occurs in the Upper Coal-Measures and in the
Transition rocks. _S. cuneifolium_[927] (Sternb.) has been recorded
from the Transition, Middle and Lower Coal-Measures. _Sphenophyllum
oblongifolium_, Germ.[928], is recorded from Lower Permian rocks, as is
also _S. Thoni_[929], Mahr.
The comparison which has naturally been drawn between _Sphenophyllum_
with its slender stems bearing occasionally dimorphic leaves, and
water-plants is not, I believe, supported by the facts of anatomy
or external characters. The entire and finely-dissected leaves
do not exhibit that regularity of relative disposition which is
characteristic of aquatic plants; the two forms of leaves may occur
indiscriminately on the same branch. The well-developed and thick xylem
is not in accordance with the anatomical features usually associated
with water-plants. It is true that in some living dicotyledons of the
family Leguminosae, which inhabit swampy places, the secondary xylem is
represented by a thick mass of unlignified and thin-walled parenchyma,
as in the genus _Aeschynomene_[930], from which the material of
‘pith’-helmets is obtained; but the wood of _Sphenophyllum_ was
obviously thick-walled and thoroughly lignified.
It is not improbable that the long and slender stems of this plant may
have grown like small lianas in the Coal-Measure forests, supporting
themselves to a large extent on the stouter branches of Calamites
and other trees. The anatomical structure of a _Sphenophyllum_ stem
would seem to be in accord with the requirements of a climbing plant.
It has been shewn[931] that in recent climbing plants the tracheae
and sieve-tubes are characterised by their large diameter, a fact
which may be correlated with the small diameter of climbing stems and
the need for rapid transport of food material. In _Sphenophyllum_
the tracheae of the xylem have a wide bore, and in _S. insigne_ the
phloem contains unusually wide sieve-tubes. The central position of
the stele is another feature which is not inconsistent with a climbing
habit. Schwendener and others[932] have demonstrated that in climbing
organs, as in underground stems and roots, there is a tendency towards
a centripetal concentration of mechanical or strengthening tissue.
The axial xylem strand of _Sphenophyllum_ would afford an efficient
resistance to the tension or pulling force which climbing stems
encounter.
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=Duval-Jouve, J.= (64) Histoire naturelle des Equisetum de France.
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=Engelhardt, H.= (87) Über _Rosselinia conjugata_ (Beck.) eine neue
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=Etheridge, R.= (81) _Vide_ Hicks, H.
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—— (95) On the occurrence of a plant in the Newcastle or Upper
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=Ettingshausen, C. von.= (55) Die Steinkohlenflora von Radnitz in
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=Ettingshausen, C. von= and =Gardner, J. S.= (79) A monograph of the
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=Feilden, H. W.= (96) Note on the glacial Geology of Arctic Europe and
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=Feistmantel, O.= (73) Das Kohlenvorkommen bei Rothwaltersdorf in
der Grafschaft Glatz und dessen organische Einschlüsse. _Zeit.
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_Palaeontograph._ vol. XXIII. p. 1, 1875–76.
—— (81) The Fossil Flora of the Gondwana System. _Mem. Geol.
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—— (90) Geological and palaeontological relations of the Coal and
plant-bearing beds of Palaeozoic and Mesozoic age in Eastern
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=Felix, J.= (86) Untersuchungen über den inneren Bau westfälischer
Carbon-Pflanzen. _Abh. Geol. Landesanst._ 1886, p. 153.
—— (94) Studien über fossile Pilze. _Zeit. deutsch. geol. Ges._ Heft
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=Fliche, P.= and =Bleicher=, —. Étude sur la flore de l’oolithe
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=Früh, J.= (90) Zur Kenntniss der gesteinbildenden Algen. _Abh.
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=Fuchs, T.= (94) Über eine fossile _Halimeda_ aus dem Eocänen Sandstein
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=Gardner, J. S.= (79) _Vide_ Ettingshausen.
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—— (45) Tchihatcheff’s Voyage Scientifique dans l’Altaï oriental,
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—— (60) Über die fossile Flora der Silurischen, der Devonischen und
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—— (64) Die fossile Flora der Permischen Formation.
_Palaeontograph._ vol. XII. 1864–65.
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—— (82) Mémoire sur la formation de la houille. _Ann. Mines_, vol.
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—— (87) Formation des couches de houille et du terrain houiller.
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=Harrison, J. B.= (91) _Vide_ Jukes-Browne.
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=Harvey, W. H.= (58) Phycologia Australica. _London_, 1858.
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=Hick, T.= (78) and (81) _Vide_ Cash, W.
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=Hooker, W. J.= (20) Musci Exotici. Vol. II. _London_, 1820.
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_Quart. Journ. Geol. Soc._ vol. XL. p. 178, 1884.
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=Huxley, T. H.= (93) Science and Hebrew tradition. _Collected Essays_,
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=James, J. F.= (93) Studies in the problematic organisms. No. 2. The
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—— (93³) Fossil fungi. Translated from the French of R. Ferry, with
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=Judeich, J. F.= and =Nitsche, F.= (95) Lehrbuch der mitteleuropäischen
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=Kent, S.= (93) The Great Barrier Reef of Australia. _London_, 1893.
=Kickx, J. J.= (64) _Vide_ Coemans, E.
=Kidston, R.= (83) Report on fossil plants collected by the Geological
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—— (83²) On the affinities of the genus _Pothocites_, Paterson; with
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—— (86) Catalogue of the Palaeozoic plants in the department of
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—— (88) _Vide_ Young.
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=Knowlton, F. H.= (89) Fossil wood and lignite of the Potomac
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—— (96) The nomenclature question. _Bot. Gazette_, vol. XXI. p. 82,
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=Koechlin-Schlumberger, J.= _Vide_ Schimper, W. P.
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=König, C.= (29) _Vide_ Murchison (29) p. 298.
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=Lesquereux, L.= (66) Geological Survey of Illinois. Vol. II. _New
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—— (70) _Ibid._ vol. IV. 1870.
—— (79) Atlas to the Coal Flora of Pennsylvania and of the
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—— (80) Description of the Coal Flora of the Carboniferous formation
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—— (84) _Ibid._ vol. III. 1884.
—— (87) A species of Fungus recently discovered in the shale of the
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=Mougeot, A.= (44) _Vide_ Schimper.
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—— (96) _Nematophyton Ortoni_, n. sp. _Annals Bot._ vol. X. p. 41,
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=Petzholdt, A.= (41) Über Calamiten und Steinkohlenbildung. _Dresden_
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=Pfitzer, E. H. H.= (67) Über die Schutzscheide. _Pringsh. Jahrb. Wiss.
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=Philippi, R. A.= (37) Beweis das die Nulliporen Pflanzen sind.
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=Phillips, J.= (29) Illustrations of the Geology of Yorkshire. _York_,
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=Phillips, W.= (93) The breaking of the Shropshire Meres. _Midland
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=Plot, R.= (1705) The Natural History of Oxfordshire. _Oxford_, 1705.
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—— (90) Der im Lichthof der Königl. Geol. Landesanstalt und
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—— (92) Der äussere Bau der Blätter von _Annularia stellata_
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—— (93) Die Flora des Rothliegenden von Thüringen. (Th. I.) _Jahrb.
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—— (94) Über die Stellung der Sphenophyllum im System. _Ber.
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—— (96) Die floristische Gliederung des deutschen Carbon und Perm.
_Abhand. k. Preuss. geol. Landesanst._ (N. F.) Heft. XXI. p. 1,
1896.
—— (96²) Palaeophytologische Notizen. _Naturwiss. Wochenschrift_,
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=Quekett, J.= (54) Lectures on Histology. Vols. I. and II. _London_,
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=Raciborski, M.= (94) Flora Kopalna ogniotrwalych Glinek Krakowskich.
_Pamiȩt. Akad. Umiejȩtności_, 1894.
=Reinsch, P. F.= (81) Neue Untersuchungen über die Mikrostruktur der
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=Renault, B.= (73) Recherches sur l’organisation des Sphenophyllum et
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—— (76) Recherches sur la fructification de quelques végétaux. _Ann.
Sci. Nat._ [6] vol. III. p. 5, 1876.
—— (76²) Nouvelles recherches sur la structure des Sphenophyllum.
_Ann. Sci. Nat._ [6] vol. IV. p. 277, 1876.
—— (82) Cours de botanique fossile. Vol. II. _Paris_, 1882.
—— (85) Recherches sur les végétaux fossiles du genre Astromyelon
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—— (93) Bassin Houiller et Permien d’Autun et d’Epinac. (_Atlas._)
_Études des gîtes minéraux de la France_, Fasc. IV. _Paris_, 1893.
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—— (95²) Sur quelques Bactéries des temps primaires. _Autun_, 1895.
—— (96) Bassin Houiller et Permien d’Autun et d’Epinac. (_Text._)
_Études des gîtes minéraux de la France_, Fasc. IV. _Paris_, 1896.
—— (96²) Notice sur les travaux scientifiques. _Autun_, 1896.
—— (96³) Recherches sur les Bactériacées fossiles. _Ann. Sci. Nat._
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=Renault, R.= and =Bertrand= (94) Sur une bactérie coprophile de
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=Renault, R.= and =Zeiller, R.= (88) Études sur le terrain houiller de
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=Reuss, A. E.= (61) Über die fossile Gattung _Acicularia_ d’Archiac.
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=Rodway, J.= (95) In the Guiana Forest. _London_, 1895.
=Römer, A.= (54) Beiträge zur geologischen Kenntniss des nordwestlichen
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=Rosanoff, S.= (66) Recherches anatomiques sur les Mélobésiées. _Mém.
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=Rosenvinge, L. K.= (93) Grønlands Havalger. Conspectus florae
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=Rothpletz, A.= (80) Die Flora und Fauna der Carbon Culmformation bei
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—— (91) Fossile Kalkalgen aus den Familien der Codiaceen und der
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—— (94) Ein geologischer Querschnitt durch die Ost-Alpen nebst
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—— (81) _Vide_ Saporta and Marion.
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—— (91) Plantes Jurassiques. Vol. IV. _Paris_, 1891.
=Saporta, de G.= and =Marion, A. F.= (81) L’Évolution du règne végétal.
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=Schenk, A.= (65) _Vide_ Schoenlein.
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=Scheuchzer, J. T.= (1723) Herbarium diluvianum. _Lugduni Batavorum_,
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=Schiffner, V.= and =Müller, C.= (95) Engler und Prantl; Die
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=Schimper, W. P.= (65) Euptychium Muscorum Neocaledonicorum genus novum
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—— (74) _Ibid._ Atlas.
=Schimper, W. P.= and =Koechlin-Schlumberger, J.= (62) Mémoire sur
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=Schimper, W. P.= and =Mougeot= (44) Monographie des plantes fossiles
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=Schimper, W. P.= and =Schenk, A.= (90) Palaeophytologie. _Zittel’s
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=Schlotheim, E. F. von.= (04) Beschreibung merkwürdiger
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=Schmitz, F.= (97) Rhodophyceae. Engler und Prantl; Die natürlichen
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=Schoenlein, J. L.= and =Schenk, A.= (65) Abbildungen von fossilen
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=Schröter, J.= (89) Engler und Prantl; Die natürlichen
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=Schulze, C. F.= (1755) Kurtze Betrachtung derer Kräuterabdrücke im
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=Schütt, F.= (93) Das Pflanzenleben der Hochsee. _Kiel_ and _Leipzig_,
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=Schwendener, S.= (74) Das mechanische Princip im anatomischen Bau der
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=Scott, D. H.= (94) (95) (96) _Vide_ Williamson, W. C.
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—— (96²) Presidential address. (Liverpool Meeting.) _British Assoc.
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=Seeman, B.= (65) A gigantic _Equisetum_. _Journ. Bot._ vol. III. p.
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=Seward, A. C.= (88) On _Calamites undulatus_. _Geol. Mag._ vol. V. p.
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—— (89) _Sphenophyllum_ as a branch of _Asterophyllites_. _Mem. and
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—— (94²) Catalogue of the Mesozoic plants in the Department of
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—— (95) _Ibid._ Pt. II. _London_, 1895.
—— (95²) Coal: its structure and formation. _Science Progress_, vol.
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—— (95³) Notes on _Pachytheca_. _Proc. Phil. Soc. Cambridge_, vol.
VIII. pt. 5, p. 278, 1895.
—— (96) Notes on the geological history of Monocotyledons. _Annals
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—— (96²) _Vide_ Feilden, H. W.
—— (97) On _Cycadeoidea gigantea_, a new Cycadean stem from the
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=Sharpe, S.= (68) On a remarkable incrustation in Northamptonshire.
_Geol. Mag._ vol. V. p. 563, 1868.
=Sherborn, C. D.= (93) An index to the genera and species of the
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=Shrubsole, W. H.= and =Kitton, F.= (81) The Diatoms of the London
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=Skertchly, J. B. J.= (77) The geology of the Fenland. _Mem. Geol.
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=Sollas, W. J.= (86) On a specimen of slate from Bray-Head traversed by
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=Solms-Laubach, Graf zu.= (81) Die corallinen Algen des Golfes von
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—— (91) Fossil Botany. _Oxford_, 1891.
—— (93) Über die Algen-genera _Cymopolia_, _Neomeris_ und
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—— (95) William Crawford Williamson. (Obit. notice.) _Nature_, vol.
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—— (95²) Über Devonische Pflanzenreste aus den Lenneschiefern der
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—— (95³) Monograph of the Acetabularieae. _Trans. Linn. Soc._ [2]
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—— (95⁴) _Bowmanites Römeri_, eine neue
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_Wien_, vol. XLV. Heft 2, p. 225, 1895.
—— (96) Über die seinerzeit von Unger beschriebenen Struktur
bietenden Pflanzenreste des Unterculm von Saalfeld in Thüringen.
_Abhand. k. Preus. Geol. Landesanst._ (N.F.) Heft XXIII.
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—— (97) Über die in den Kalksteinen des Culm von Glätzisch
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=Sorby, H. C.= (79) Presidential Address. _Proc. Geol. Soc._ 1878–79,
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=Sprengel, A.= (28) Commentatio de Psarolithis. _Halle_, 1828.
=Stefani, K. de.= (82) Vorläufige Mittheilung über die rhätischen
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=Steinmann, G.= (80) Zur Kenntniss fossiler Kalkalgen (Siphoneae).
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=Stenzel, G.= (86) _Rhizodendron oppoliense_, Göpp. _Jahresbericht.
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=Sternberg, C. von.= (20) Versuch einer geognostisch-botanischen
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—— (21) _Ibid._ Fasc. II.
—— (25) _Ibid._ Fasc. IV.
—— (38) _Ibid._ Fasc. VIII.
=Sterzel, J. T.= (82) Über die Fruchtähren von _Annularia
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INDEX.
_Acetabularia_, 165, 166
_A. mediterranea_, 162, 165, 166
_Achyla_, 127
_Acicularia_, 162, 166, 167
_A. Andrussowi_, 162, 168
_A. miocenica_, 162, 168
_A. pavantina_, 167
_A. Schencki_, 162, 166, 168
_Aegoceras planicosta_, 61
Aeolian rocks, 25
_Aeschynomene_, 414
Africa, South, 80, 284
_Agaricus melleus_, 211, 212, 215
Algae, 138–202
_Algites_, 148, 149, 204, 205
Amazons, 66
Amber, 70, 71, 206, 211, 221, 255
America, 182
_Amphiroa_, 184
_Anarthrocanna_, 283
_A. tuberculata_, 386
Andersson, G., 60
Andrae, K. J., 276
Andreaeales, 236
Andrussow, N., 168
_Annularia_, 255, 260, 282, 283, 289, 329, 332, 333, 336–339, 340,
361, 362, 379, 381
_A. calamitoides_, 335
_A. Geinitzi_, 338
_A. longifolia_, 338, 341
_A. radiata_, 283
_A. ramosa_, 363
_A. sphenophylloides_, 340, 341, 363
_A. stellata_, 338–341, 363
_A. westphalica_, 338
Antigua, 79
_Aphlebia_, 142
_Araucaria imbricata_, 101, 104
_Araucarioxylon Withami_, 81, 82
Archaean system, 34–36
_Archaeocalamites_, 254, 255, 256, 263, 285, 336, 383–388
_A. scrobiculatus_, 385–387
d’Archiac, A., 166, 167
Arctic plants, 16, 17
Arenig series, 37
Arran, 88, 89
_Arthrodendron_, 301, 302, 324, 325, 326, 327, 366, 379, 381
_Arthropitys_, 300, 301, 302, 304, 311, 324, 325, 326, 328, 333, 349,
369, 375, 379, 380, 381, 384
_A. approximata_, 371
_A. bistriata_, 326
Arthropod eggs, 108
Artis, F. T., 4, 104, 297
_Artisia_, 104
Ascomycetes, 208, 209, 220
Asia Minor, Coal-Measures of, 282, 283
_Asterocalamites scrobiculatus_, 386
_Asterophyllites_, 264, 276, 329, 332, 333, 338, 369, 371, 379, 397,
409
_A. elegans_, 386
_A. equisetiformis_, 335, 338
_A. insignis_, 397
_A. longifolius_, 338
_A. spaniophyllus_, 386
_A. sphenophylloides_, 397
_Astromyelon_, 342, 343, 381
Australia, 179, 290, 291
Autun, 6, 46, 83, 179, 181, 182, 206, 390, 399
Azoic system, 36
Bacillariaceae, 150–156
_Bacillus_, 135, 136
_B. amylobacter_, 136
_B. permicus_, 135
_B. Tieghemi_, 135, 136
Bacteria, 75, 132, 138
_Bactryllium_, 154–156
_B. deplanatum_, 155
Bala series, 37
_Balanoglossus_, 145
Balfour, Bayley, 178
Balfour, J. H., 101
_Bambusa arundinacea_, 322
Barbados, 188
Barber, C. A., 193, 197, 198, 203, 204
Barrois, C., 146
Barton, R., 80
de Bary, A., 132
Basidiomycetes, 208, 211, 212
Bates, H. W., 64, 66
Bauhin, C., 225
Bear Island, 366
Bentham, G., 98, 99
_Bergeria_, 101
Bernician series, 43
Bertrand, C. E., 87, 154, 178, 181
Beyrich, E., 52
Bilin, 152
_Bilobites_, 148
Binney, E. W., 9, 10, 304, 311, 343, 351, 386, 401
Bituminous deposits, 178–183
Blackman, V. H., 119, 120
_Blandowia_, 231
Boghead, 178–183
Boring algae, 127, 128
Boring fungi, 127, 128
Bornemann, J. G., 129, 130, 148, 177
Bornet, E., 181
Bornet, E. and Flahault, 128, 129
_Bornetella_, 177
_Bornia_, 263
_B. equisetiformis_, 335
_B. scrobiculata_, 386
_B. stellata_, 338
_Bostrychus_, 210, 211
_Bothrodendron_, 133, 134
_Botrydium_, 157
_Bowmanites_, 401, 402
_B. Dawsoni_, 401
_B. Germanicus_, 408
Brazil, 79
Brodie, P. B., 240
Brongniart, A., 5, 6, 9, 11, 17, 18, 111, 112, 145, 177, 233, 269,
281, 288, 289, 291, 297, 299, 300, 301, 332, 336, 351, 390, 391
Bronn, H. G., 118
Brora, 269
Brown, A., 189, 190
_Brukmannia_, 350, 360, 361
Bryales, 236
Bryophyta, 229–241, 243
Bryozoa, 187
_Buccinum_, 159
Buckman, J., 240, 278
Bunbury, Sir C. J. F., 270, 271, 273, 284, 287, 288, 289
Bunter series, 47, 267, 292, 293
Burntisland, 88, 90, 397
_Buthotrephis radiata_, 338
Büzistock, 28
Calamarieae, 255, 295–388
_Calamitea_, 298, 300, 301, 303
_C. bistriata_, 301
_Calamites_, 18, 19, 77, 86, 94, 104, 114, 252, 254–257, 259, 260,
262, 264, 269, 285, 286, 289, 295–383, 386–388, 413
_Calamites_, sub-genera of, 381
_C. alternans_, 378
_C. approximatus_, 369, 370, 374, 378
_C. arborescens_, 363
_C. Beani_, 270, 273
_C. cannaeformis_, 374
_C. communis_, 326, 351
_C. cruciatus_, 376–378
_C. decoratus_, 374
_C. equisetiformis_, 335
_C. giganteus_, 271
_C. gigas_, 271
_C. Göpperti_, 368, 372–374
_C. lateralis_, 276
_C. laticulatus_, 386
_C. obliquus_, 386
_C. pedunculatus_, 358
_C. ramosus_, 363
_C. Sachsei_, 372
_C. scrobiculatus_, 386
_C. Suckowi_, 372, 374, 375
_C. transitionis_, 386
_C. varians_, 371
_C. variolatus_, 386
_C. verticillatus_, 372
_Calamitina_, 260, 266, 267, 329, 330, 362, 364, 365, 367–376, 381,
383, 388
_C. pauciramis_, 367
_Calamocladus_, 227, 289, 328, 329, 332, 333, 336, 337, 350, 361–364,
381, 409
_C. equisetiformis_, 333–335, 363
_C. frondosus_, 289
_Calamodendron_, 300–302, 328, 349, 364, 375, 378, 379, 380, 382
_C. bistriatum_, 300
_C. commune_, 311, 317
_C. cruciatum_, 378
_C. intermedium_, 328
_C. striatum_, 300
_Calamophyllites_, 371
_C. Göpperti_, 372
_Calamopitys_, 301, 302, 381
_Calamostachys_, 339, 342, 350, 351–357, 362, 371, 379, 381, 384,
387, 402
_C. Binneyana_, 351–355, 356, 357, 361
_C. calathifera_, 341, 363
_C. Casheana_, 355–357, 361
_C. longifolia_, 363
_C. ramosa_, 363
_C. Solmsi_, 363, 364
_C. tenuissima_, 261
_C. tuberculata_, 363
Calciferous sandstone, 43, 148, 397, 412
Callus wood, in _Calamites_, 319, 320
_Calothrix_, 126
Cambrian, 36, 37, 149, 177
de Candolle, A., 297
Carboniferous limestone, 40, 42, 43, 387
Carboniferous system, 39–45
Carinal canals, 249, 306, 307, 308
Carlsbad, 123, 126
Carpenter, W. B., 167
_Carpolithus_, 280
Carruthers, W., 9, 10, 87, 193, 197, 202, 212, 217, 240, 304, 351
Cash, W. and Hick, T., 206, 218, 219, 342
Castracane, F., 154
_Casuarina stricta_, 95, 96
_Casuarinites_, 332
_C. equisetiformis_, 335
_C. stellata_, 338
Caterpillars, Liassic, 240
_Caulerpa_, 157–159
_C. abies-marina_, 142
_C. cactoides_, 142, 158
_C. Carruthersi_, 159
_C. ericifolia_, 142
_C. plumaris_, 142
_C. pusilla_, 142
_C. scalpelliformis_, 142
_C. taxifolia_, 142
Caulerpaceae, 157–159
_Caulerpites_, 142, 158
_C. cactoides_, 158
_Celastrum_, 181
_Celluloxylon primaevum_, 198
_Ceratium_, 117
Chalk, 26, 50, 120
Challenger, H. M. S., 65, 66, 101, 117, 118, 119, 122, 151
_Chara_, 69, 223, 228
_C. Bleicheri_, 226
_C. foetida_, 224, 225
_C. Jaccardi_, 226
_C. Knowltoni_, 224, 226, 227
_C. Wrighti_, 226, 227
Characeæ, 222–228
Chareae, 223–228
Charophyta, 222–228
_Cheirostrobus_, 258, 413
_C. Pettycurensis_, 412
Chert, 227
Chlorophyceae, 127–130, 156–190
_Chondrites_, 142, 144
_C. plumosa_, 148
_C. verisimilis_, 146, 147
_Chondrus_, 231
_C. crispus_, 191
Chroococcaceae, 122, 123, 181
Church, A. H., 171
Chytridineae, 216
_Cingularia_, 290, 364
Cladochytrium, 216
_Cladosporites bipartitus_, 217, 220
Clayton, fossil tree, 71
Coal, 44, 45, 68, 76, 92, 133, 134, 178, 179
Coal-balls, 85, 86
Coalbrook Dale, 361
Coal-Measures, 40–45, 55, 84, 85, 217, 238, 256, 259, 261, 268,
282, 283, 296, 301, 302, 325, 329, 358, 375, 376, 378, 382, 397,
401, 406, 413
Coccoliths, 120
Coccospheres, 118–121
Codiaceae, 159–164
_Codium_, 159, 160
_C. Bursa_, 160
_C. tomentosum_, 160
_Coelotrochium_, 176
Coemans, E. and Kickx, J. J., 411
Coenocyte, 116, 117
Cohn, F., 123, 126
Cole, G., 83
Cone-in-cone structure, 83
_Confervites_, 177
_C. chantransioides_, 178
Confervoideae, 177–178
Conglomerate, 24
Coniferae, 232
_Convallarites_, 291
Conwentz, H., 80, 211, 212, 221
Cook, Capt., 122
Coprolites, 108, 135, 137, 182
Coral reefs, 26, 40, 48
_Corallina officinalis_, 183, 184
Corallinaceae, 183–190
Corallineae, 184
Corallines, 159, 169, 184
_Coralliodendron_, 163
Corals, fossil, 25
Corda, A. J., 5, 8
_Cordaites_, 75, 80, 99, 104, 366
Cormack, B. G., 251, 252
_Coscinodiscus_, 153
Costa, Mendes da, 3
Cotta, C. B., 9, 298
‘Crag’, 53, 62
Craigleith Quarry, Edinburgh, 81
Cramer, C., 171, 177
Credner, H., 149
Crépin, F., 109
Cretaceous system, 50, 51, 188
Cromer Forest bed, 53
_Crossochorda_, 148
_Cruziana_, 144, 145
Culm rocks, 42, 68, 383, 413
Cuticles, fossil, 68, 133
Cyanophyceae, 121, 122–132
_Cyathophyllum bulbosum_, 237
Cycadaceæ, 98
_Cycadeoidea gigantea_, 88, 214
_Cycadorachis_, 114, 115
Cycads, 49, 56, 99, 281
_Cyclocladia_, 371
_C. major_, 372
_Cymopolia_, 165, 169, 170, 172, 177
_C. barbata_, 162, 169, 171
_C. elongata_, 172
_C. Mexicana_, 169
_Dactylopora_, 175, 176
_D. cylindracea_, 176
_Dactyloporella_, 176
Darwin, C., 16, 17, 65, 70, 79, 100
Dasycladaceae, 164–176
_Dasyporella_, 176
Dawes, J. S., 297, 302
Dawson, Sir W., 147, 192, 193, 195, 338, 390, 413
_Dawsonia_, 237
_D. polytrichoides_, 237
_D. superba_, 237
Decorticated stems, 105
Deecke, W., 173
Defrance, J. L. M., 172
_Dendrophycus triassicus_, 146
Denudation, 23, 24, 35
Desmideae, 221
Devonian system, 39, 133, 173, 193, 195, 225, 256, 338, 383, 413
Diatomaceae, 150–156
Diatomite, 151
Diatoms, 117, 118, 123, 141, 185
_Dichothrix_, 123
Dicotyledons, 98, 99
Dictyogens, 99
Dinoflagellata, 118
_Diplopora_, 173–176
Dirt-beds, of Portland, 56, 57
Dismal swamp, 74
Dixon, H. N. and Joly, J., 120
Dorset, coast, 56, 226
Dover coal, 45
Drifting of trees &c., 64
Duncan, M., 127, 129
Duval-jouve, J., 247
Dyas system, 46, 209
Echinoid spines, 177
Ehrenberg, C. G., 117, 118, 151, 153
Elaters, 245
Ellis, J., 159, 169
Endophytic algae, 132
Eocene rocks, 29, 52, 164, 171
_Eophyton_, 144, 148
_Eopteris Morierei_, 106
_Ephedra distachya_, 95, 97
Equisetaceae, 19, 244–254, 255, 256
Equisetales, fossil, 254–388
Equisetales, recent, 244–254
_Equisetites_, 254, 257–273, 282, 291, 330, 413
_E. arenaceus_, 268, 269, 275, 280
_E. Beani_, 270–275
_E. Brodii_, 278
_E. Burchardti_, 279, 280
_E. Burejensis_, 280
_E. columnaris_, 72, 265, 269–271
_E. Göpperti_, 386
_E. Hemingwayi_, 259, 262–264
_E. lateralis_, 265, 275–279
_E. Monyi_, 266
_E. Mougeoti_, 267
_E. Münsteri_, 278, 279
_E. Parlatori_, 280
_E. platyodon_, 267
_E. rotiferum_, 279
_E. spatulatus_, 264, 265
_E. Vaujolyi_, 261
_E. Yokoyamae_, 280
_E. zeaeformis_, 265–266
_Equisetum_, 19, 245–254, 258, 262, 263, 267, 268, 281, 282, 285,
286, 287, 290, 292, 297, 298, 304, 307, 308, 309, 327, 337, 346,
360, 381, 384, 388, 390
_E. arvense_, 246, 247, 251, 279
_E. debile_, 95, 96, 97
_E. giganteum_, 245, 297
_E. hiemale_, 270
_E. limosum_, 263
_E. litorale_, 253
_E. maximum_, 245, 246, 247, 250–253
_E. palustre_, 247, 253
_E. ramosissimum_, 259, 265, 270
_E. sylvaticum_, 247, 279
_E. Telmateia_, 245
_E. trachyodon_, 270
_E. variegatum_, 95, 252
_E. xylochaetum_, 250
_Eryngium Lassauxi_, 99
_E. montanum_, 99
Etheridge, R., 200, 215, 290
Ettingshausen, C. von, 6, 7, 311, 332, 372
_Eucalamites_, 367, 376, 379
_Eurypterus_, 371
_E. mammatus_, 371
Feilden, Col. H. W., 211
Feistmantel, O., 284, 291, 293, 387
Felix, J., 399
Fischer, A., 217
Fish-scales, 127, 182
_Fleurs d’eau_, 132, 182
Fliche, P. and Bleicher, 232
Flint, 62
Florideae, 175, 183–190
Flowers, fossil, 70
Flysch, 147, 148, 192
_Fontinalis_, 240
Foraminifera, 26, 161, 163, 175, 185
Forbes, E., 16
Forbes, H. O., 63
Forest-bed, 53