affinities to any members of the group, and the view, of which Williamson was the chief advocate, that they form a homogeneous Cryptogamic family, is now fully established.
II. Sphenophyllales.—The class of Sphenophyllales, as known to us at present, is of limited extent, embracing the two genera Sphenophyllum and Cheirostrobus, which may serve as types of two families within the class. The characters of Sphenophyllum are known with some completeness, while our knowledge of Cheirostrobus is confined to the fructification; the former will therefore be described first.
1. Sphenophyllum.—The genus Sphenophyllum, of which a number of species have been described, ranging probably from the Middle Devonian, through the Carboniferous, to the Permian or even the Lower Triassic, consisted of herbaceous plants of moderate dimensions. The long, slender stems, somewhat tumid at the nodes, were ribbed, the ribs running continuously through the nodes, a fact correlated with the superposition of the whorled leaves, the number of which in each verticil was some multiple of 3, and usually 6. In the species on which the genus was founded the leaves, as the generic name implies, are cuneate and entire, or toothed on their anterior margin;[1] in other cases they are deeply divided by dichotomy into narrow segments, or the whorl consists of a larger number (up to 30) of apparently simple, linear leaves, which may represent the segments of a smaller number. The different forms of leaf may occur on the same plant, the deeply divided foliage often characterizing the main stem, while the cuneate leaves were borne on lateral shoots. A comparison, formerly suggested, with the two forms of leaf in Batrachian Ranunculi has not proved to hold good; the idea of an aquatic habit is contradicted by the anatomical structure, and the hypothesis that the plants were of scandent growth is more probable. The species of Sphenophyllum have a graceful appearance, which has been compared with that of the trailing Galiums of hedgerows. Branches sprang from the nodes, though perhaps not truly axillary in position. The cones, more or less sharply differentiated, terminated certain of the branches.
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| Fig. 6.—Sphenophyllum Dawsoni. Diagram of cone in longitudinal section. | |
| 𝑎𝑥, | Axis. |
| 𝑏𝑟, | Bracts. |
| 𝑠𝑝, | Sporangiophores, each bearing a sporangium, 𝑠𝑚. |
| 𝑏𝑟′, | Whorl of bracts in surface view. |
The anatomy of the stem of Sphenophyllum, investigated by Renault, Williamson and others, is highly characteristic (fig. 5, Plate). The stem is traversed by a single stele, with solid wood, without pith; the primary xylem is triangular in section, the spiral elements forming one or two groups at each angle, while the phloem occupied the bays, so that the structure resembles that of a triarch root. Two leaf-trace bundles started from each angle of the stele, and forked, in passing through the cortex, to supply the veins of the leaf, or its subdivisions. The cortex was deeply furrowed on its outer surface. The primary structure is only found unaltered in the youngest stems; secondary growth by means of a cambium set in very early, xylem being formed internally and phloem externally in a perfectly normal manner. At the same time a deep-seated periderm arose, by which the primary cortex was soon entirely cut off. The secondary wood in the Lower Carboniferous species, S. insigne, has scalariform tracheides, and is traversed by regular medullary rays, but in the forms from later horizons the tracheides are reticulately pitted, and the rays are for the most part replaced by a network of xylem-parenchyma. There are no recent stems with a structure quite like that of Sphenophyllum; so far as the primary structure is concerned, the nearest approach is among the Psiloteae, with which other characters indicate some affinity; the base of the stem in Psilotum forms some secondary wood. The diarch roots of a Sphenophyllum have been described by Renault, who has also investigated the leaves; they were strongly constructed mechanically, and traversed by slender vascular bundles branching dichotomously.
Fructification.—Williamson thoroughly worked out, in petrified specimens, the organization of a cone which he named Bowmanites Dawsoni; it was subsequently demonstrated by Zeiller that this fructification belonged to a Sphenophyllum, the cones of the well known species S. cuneifolium having a practically identical structure. The type of fructification described by Williamson and now named Sphenophyllum Dawsoni consists of long cylindrical cones, in external habit not unlike those of some Calamarieae. The axis, which in structure resembles the vegetative stem in its primary condition, bears numerous verticils of bracts, those of each verticil being coherent in their lower part, so as to form a disc or cup, from the margin of which the free limbs of the bracts arise. The sporangia, which are about twice as numerous as the bracts, are seated singly on pedicels or sporangiophores springing from the upper surface of the bract-verticil, near its insertion on the axis (fig. 6). As a rule two sporangiophores belong to each bract. The sporangium is attached to the enlarged distal end of its pedicel, from which it hangs down, so as to suggest an anatropous ovule on its funiculus. Dehiscence appears to have taken place at the free end of the sporangium; the spores are numerous, and, so far as observed, of one kind only. Each sporangiophore is traversed throughout its length by a vascular bundle connected with that which supplies the subtending bract. This form of fructification appears, from Zeiller’s researches, to have been common to several species of Sphenophyllum, but others show important differences. Thus Bowmanites Römeri, a fructification fully investigated by Solms-Laubach, differs from S. Dawsoni in the fact that each sporangiophore bears two sporangia, attached to a distal expansion approaching the peltate scale of the Equisetales. It is thus proved that the sporangiophore is not a mere sporangial stalk, but a distinct organ, in all probability representing a ventral lobe of the subtending bract. The recently discovered species, Sphenophyllum fertile, while resembling Bowmanites Römeri in its peltate, bisporangiate sporangiophores, is peculiar in the fact that both dorsal and ventral lobes of the sporophyll were fertile, dividing in a palmate manner into several branches, each of which constitutes a sporangiophore. Thus the sterile bracts of other species are here replaced by sporangium-bearing organs. In Sphenophyllum majus, where the cones are less sharply defined, the forked bract bears a group of four sporangia at the bifurcations, but their mode of insertion has not yet been made out.
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Fig. 7.—Cheirostrobus. Diagram of cone, the upper part in transverse, the lower in longitudinal section. In the transverse section six sporophylls, each showing three segments, are represented. | |
| S𝑝.𝑎, | Section through sterile segments. |
| S𝑝.𝑏, | Section through sporangiophores. |
| 𝑠𝑡, | Laminae of sterile segments. |
| 𝑓, | Peltate expansions of sporangiophores. |
| 𝑠𝑚, | Sporangia. |
| 𝑣.𝑏. | Vascular bundles. |
| 𝑐𝑦, | Stele of axis (A𝑥). |
| In the longitudinal section the corresponding parts are shown. | |
2. Cheirostrobeae.—The family Cheirostrobeae is only known from the petrified fructification (Cheirostrobus pettycurensis) derived from the Lower Carboniferous of Burntisland in Scotland. The excellence of the preservation of the specimens has rendered it possible to investigate the complex structure in detail. The cone is of large size—3·5 cm. in diameter; the stout axis bears numerous whorls of compound sporophylls, the members of successive verticils being superposed. The sporophylls, of which there are eleven or twelve in a whorl, are each composed of six segments, three being inferior or dorsal, and three superior or ventral. The dorsal segments are sterile, corresponding to the bracts of Sphenophyllum Dawsoni, while the ventral segments constitute peltate sporangiophores, each bearing four sporangia, just as in a
- ↑ In S. speciosum the leaves in a whorl were of unequal size.
