Calamarian fructification (fig. 7). The great length and slender
proportions of the segments give the cone a peculiar character,
but the relations of position appear to leave no doubt as to the
homologies with the fructification of Sphenophylleae; as regards
the sporangiophores, Bowmanites Römeri occupies exactly the
middle place between S. Dawsoni and Cheirostrobus. The axis of
the cone in Cheirostrobus contains a polyarch stele, with solid
wood, from the angles of which vascular bundles pass out, dividing
in the cortex, to supply the various segments of the sporophylls.
In the peduncle of the strobilus secondary tissues are formed.
While the anatomy has a somewhat Lycopodiaceous character,
the arrangement of the appendages is altogether that of the Sphenophylleae;
at the same time Calamarian affinities are indicated by
the characters of the sporangiophores and sporangia.
The Sphenophyllales as a whole are best regarded as a synthetic group, combining certain characters of the Ferns and Lycopods with those of the Equisetales, while showing marked peculiarities of their own. Among existing plants their nearest affinities would appear to be with Psiloteae, as indicated not merely by the anatomy, but much more strongly by the way in which the sporangia are borne. There is good reason to believe that the ventral synangium of the Psiloteae corresponds to the ventral sporangiophore with its sporangia in the Sphenophyllales. Professor Thomas of Auckland, New Zealand, has brought forward some interesting variations in Tmesipteris which appear to afford additional support to this view.
Pseudobornia.—Professor Nathorst has described a remarkable Devonian plant, Pseudobornia ursina (from Bear Island, in the Arctic Ocean), which shows affinity both with the Equisetales and Sphenophyllales. The stem is articulated and branched, attaining a diameter of about 10 cm. The smaller branches bear the whorled leaves, probably four in each verticil. The leaves are highly compound, dividing dichotomously into several leaflets, each of which is deeply pinnatifid, with fine segments. When found detached these leaves were taken for the fronds of a Fern. The fructification consists of long, lax spikes, with whorled sporophylls; indications of megaspores have been detected in the sporangia. The discoverer makes this plant the type of a new class, the Pseudoborniales. At present only the external characters are known.
III. Lycopodiales.—In Palaeozoic ages the Lycopods formed one of the dominant groups of plants, remarkable alike for the number of species and for the great stature which many of them attained. The best known of the Palaeozoic Lycopods were trees, reaching 100 ft. or more in height, but side by side with these gigantic representatives of the class, small herbaceous Club-mosses, resembling those of the present day, also occurred. Broadly speaking, the Palaeozoic Lycopods, whatever their dimensions, show a general agreement in habit and structure with our living forms, though often attaining a much higher grade of organization. We will first take the arborescent Lycopods, as in every respect the more important group. They may all be classed under the one family Lepidodendreae, which is here taken to include Sigillaria.
| |
| (After Stur. Scott, Studies.) | |
| Fig. 8.—Leaf-base of a Lepidodendron. | |
| s.c., | Scar left by the leaf. |
| v.b., | Print of vascular bundle. |
| p, p, | Parichnos. |
| l, | Ligule. |
| a, a, | Superficial prints below scar. |
Lepidodendreae.—The genus Lepidodendron, with very numerous species, ranging from the Devonian to the Permian, consisted of trees, with a tall upright shaft, bearing a dense crown of dichotomous branches, clothed with simple narrow leaves, ranged in some complex spiral phyllotaxis. In some cases the foliage is preserved in situ; more often, however, especially in the main stem and larger branches, the leaves had been shed, leaving behind them their scars and persistent bases, on which the characteristic sculpturing of the Lepidodendroid surface depends. The cones, often of large size, were either terminal on the smaller twigs, or, it is alleged, borne laterally on special branches of considerable dimensions. At its base the main stem terminated in dichotomous roots or rhizophores, bearing numerous rootlets. To these underground organs the name Stigmaria is applied; they are not clearly distinguishable from the corresponding parts of Sigillaria. The numerous described species of Lepidodendron are founded on the peculiarities of the leaf-cushions and scars, as shown on casts or impressions of the stem. The usually crowded leaf-cushions are spirally arranged, and present no obvious orthostichies, thus differing from those of Sigillaria. Each leaf-cushion is slightly prominent; towards its upper end is the diamond-shaped or triangular scar left by the fall of the actual leaf (fig. 8). On the scar are three prints, the central one alone representing the vascular bundle, while the lateral prints (parichnos) mark the position of merely parenchymatous strands. In the median line, immediately above the leaf-scar, is a print representing the ligule, or rather the pit in which it was seated. On the flanks of the cushion, below the scar, are two superficial prints, perhaps comparable to lenticels. In the genus Lepidophloios the leaf-cushions are more prominent than in Lepidodendron, and their greatest diameter is in the transverse direction; on the older stems the leaf-scar lies towards the lower side of the cushion. The genus Bothrodendron, going back to the Upper Devonian, differs from Lepidodendron in its minute leaf-scars and the absence of leaf cushions, the scars being flush with the smooth surface of the stem. In the Lower Carboniferous of central Russia beds of coal occur consisting of the cuticles of a Bothrodendron, which are not fossilized, but retain the consistency and chemical composition of similar tissues in recent plants.
| |
| (Scott, Studies.) | |
| Fig. 9.—Lepidodendron Veltheimianum. Transverse section of stem. | |
| p, | Pith, almost destroyed. |
| x, | Zone of primary wood. |
| px, | Protoxylem. |
| x2, | Secondary wood. |
| ph, | Phloem and pericycle. |
| br, | Stele of a branch. |
| pd, | Periderm. |
| l.b, | Leaf-bases. |
| The primary cortex between stele and periderm has perished. (×4½.) | |
The anatomy of Lepidodendron and its immediate allies is now well known in a number of species; the Carboniferous rocks of Great Britain are especially rich in petrified specimens, which formed the subject of Williamson's extensive investigations. The stem is in all cases monostelic; in most of the forms the central cylinder underwent secondary growth, and the distinction between primary and secondary wood is very sharply marked. In L. Harcourtii, however, the species earliest investigated (by Witham, 1833, and Brongniart, 1837), and in one or two other species, no secondary wood has yet been found. The primary wood of Lepidodendron forms a continuous cylinder, not broken up into distinct bundles; its development was clearly centripetal, the spiral elements forming more or less prominent peripheral groups. In the larger stems of most species there was a central pith, but in certain of the smaller branches, and throughout the stem in some species (L. rhodumnense, L. selaginoides), the wood was solid. A single leaf-trace, usually collateral in structure, passed out into each leaf. The primary structure of the stem was thus of a simple Lycopodiaceous type, resembling on a larger scale what we find in the upright stem of Selaginella spinosa. In most species (e.g. L. selaginoides, L. Wunschianum, L. Veltheimianum) secondary growth in thickness took place, and secondary wood was added, in the centrifugal direction, showing a regular radial arrangement, with medullary rays between the series of tracheides (fig. 9). The tissue thus formed often attained a considerable thickness. While
