Ancient Plants – Past Histories of Plant Families:The Lycopods

CHAPTER XIV
PAST HISTORIES OF PLANT FAMILIES VII. The Lycopods

The present-day members of this family are not at all impressive, and in their lowliness may well be overlooked by one who is not interested in unpretending plants. The fresh green mosslike Selaginella grown by florists as ornamental borders in greenhouses and the creeping “club moss” twining among the heather on a Highland moor are probably the best known of the living representatives of the Lycopods. In the past the group held a very different position, and in the distant era of the Coal Measures it held a dominant one. Many of the giants of the forest belonged to the family, and the number of species it contained was very great.

Let us turn at once to this halcyon period of the group. The history of the times intervening between it and the present is but the tale of the dying out of the large species, and the gradual shrinking of the family and dwarfing of its representative genera.

It is difficult to give the characters of a scientific family in a few simple words; but perhaps we may describe the living Lycopods as plants with creeping stems which divide and subdivide into two with great regularity, and which bear large numbers of very small pointed leaves closely arranged round the stem. The fruiting organs come at the tips of the branches, and sometimes themselves divide into two, and in these cone-like axes the spore cases are arranged, a single one on the upper side of each of the scales. In the Lycopods the spores are all alike, in the Selaginellas there are larger spores borne in a small number (four) in some sporangia, and others in large numbers and of smaller size on the scales above them. The stems are all very slender, and have no zones of secondary wood. They generally creep or climb, and from them are put out long structures something like roots in appearance, which are specially modified stem-like organs giving rise to roots.

From the fossils of the Coal Measures Lepidodendron must be chosen as the example for comparison. The different species of this genus are very numerous, and the various fossilized remains of it are among the commonest and best known of palæontological specimens. The huge stems are objects of public interest, and have been preserved in the Victoria Park in Glasgow in their original position in the rocks, apparently as they grew with their spreading rootlike organs running horizontally. A great stump is also preserved in the Manchester Museum, and is figured in the frontispiece. While among the casts and impressions the leaf bases of the plant are among the best preserved and the most beautiful. The cone has already been illustrated, and is one of the best known of fossil fructifications.

fig93

Fig. 93.:Photo of Leaf Bases of Lepidodendron

C, Scar of leaf; S, leaf base. In the scar: v, mark of severed vascular bundle, and p, of parichnos. l, Ligule scar.

From the abundant, though scattered material, fossil botanists have reconstructed the plants in all their detail. The trunks were lofty and of great thickness, bearing towards the apex a much-branched crown, the branches, even down to the finest twigs, all dividing into two equal parts. The leaves, as would be expected from the great size of the plants, were much bigger than those of the recent species, but they were of the same relatively small size as compared with the stems, and of the same simple pointed shape. A transverse section across the apex of a fertile branch shows these closely packed leaves arranged in series round the axis, those towards the outside show the central vascular strand which runs through each.

fig94

Fig. 94.:Section across an Axis surrounded by many Leaves, which shows their simple shape and single central vascular bundle v

The markings left on the well-preserved leaf-scars indicate the main features of the internal anatomy of the leaves. They had a single central vascular strand (v, fig. 93), on either side of which ran a strand of soft tissue p called the parichnos, which is characteristic of the plants of this group. While another similarly obscure structure associated with the leaf is the little scale-like ligule l on its upper surface.

The anatomy of the stems is interesting, for in the different species different stages of advance are to be found, from the simple solid protostele with a uniform mass of wood to hollow ring steles with a pith. An interesting intermediate stage between these two is found in Lepidodendron selaginoides, where the central cells of the wood are not true water-conducting cells, but short irregular water-storage tracheides, which are mixed with parenchyma. All the genera of these fossils have a single central stele, round which it is usual to find a zone of secondary wood of greater or less extent according to the age of the plant.

fig95

Fig. 95.:Transverse Section of Lepidodendron selaginoides, showing the circular mass of primary wood, the central cells of which are irregular water-storage tracheides

s, Zone of secondary wood; c, inner cortical tissues; r, intrusive burrowing rootlet; oc, outer cortical tissues with corky external layers k. (Microphoto.)

Some stems instead of this compact central stele have a ring of wood with an extensive pith. Such a type is illustrated in fig. 96, which shows but a part of the circle of wood, and the zone of the secondary wood outside it, which greatly exceeds the primary mass in thickness. This zone of secondary wood became very extensive in old stems, for, as will be imagined, the primary wood was not sufficient to supply the large trunks. The method of its development from a normal cambium in radiating rows of uniform tracheides is quite similar to that which is found in the pines to-day. This is the most important difference between the living and the fossil stems of the family, for no living plants of the family have such secondary wood. On the other hand, the individual elements of this wood are different from those of the higher families hitherto considered, and have narrow slit-like pits separated by bands of thickening on the longitudinal walls. Such tracheides are found commonly in the Pteridophytes, both living and fossil. Their type is seen in fig. 96, B, which should be compared with that in figs. 78, A and 62, B to see the contrast with the higher groups.

fig96

fig96b

Fig. 96.:A, Lepidodendron Stem with Hollow Ring of Wood W and Zone of Secondary Wood S. B, Longitudinal View of the Narrow Pits of the Wood Elements.

To supply the vascular tissues of the leaf traces, simple strands come off from the outer part of the primary wood, where groups of small-celled protoxylem project. The leaf strands lt move out through the cortex in considerable numbers to supply the many leaves, into each of which a single one enters.

fig97

Fig. 97.:Transverse Section of Outer Part of Primary Wood of Lepidodendron, showing px, projecting protoxylem groups; lt, leaf trace coming from the stele and passing (as lt1) through the cortex

As regards the fructifications of Lepidodendron much could be said were there space. The many genera of Lepidodendron bore several distinct types of cones of different degrees of complexity. In several of the genera the cones were simple in organization, directly comparable with those of the living Lycopods, though on a much larger scale. In some the spores were uniform, all developing equally in numerous tetrads. The sporophyll was radially extended, and along it the large sausage-shaped sporangia were attached. The tips of the sporophylls overlapped and afforded protection to the sporangia. The axis of the cone had a central stele with wood elements like those in the stem. The appearance of a transverse section of an actual cone is shown in fig. 99. Here the sporangia are irregular in shape, owing to their contraction after ripeness and during fossilization. Other cones had sporangia similar in size and shape, but which produced spores of two kinds, large ones resulting from the ripening of only two or three tetrads in the lower sporangia, and numerous small ones in the sporangia above.

fig98

Fig. 98.:Longitudinal Diagram, showing the arrangement of the elongated sporangia on the sporophylls

a, Main axis, round which the sporophylls are inserted; S, sporangium; s, leaflike end of sporophyll.

The similarity between the Lepidodendron and the modern Lycopod cone has been pointed out already, and it is this which forms the principal guarantee that they belong to the same family, though the size and wood development of the palæozoic and the modern plants differ so greatly.

The large group of the Lepidodendra included some members whose fructifications had advanced so far beyond the simple sporangial cones described above as to approach very closely to seeds in their construction. This type was described on fig. 54, in a series of female fructifications, so that its essential structure need not be recapitulated.

fig99

Fig. 99.:Transverse Section through Cone of Lepidodendron

A, Main axis with woody tissue; st, stalks of sporophylls cut in oblique longitudinal direction; s, tips of sporophylls cut across; S, sporangia with a few groups of spores. (Microphoto.)

The section shown in fig. 100 is that cut at right angles to that in which the sporangia are shown in fig. 98, viz. tangential to the axis. A remarkable feature of the plant is that there were also round those sporangia which bore the numerous small spores (corresponding to pollen grains) enclosing integument-like flaps similar to those shown in fig. 100, sp. f.

fig100

Fig. 100.:Section through one Sporangium of Lepidocarpon

sp, Sporophyll; sp.f., flaps of sporophyll protecting sporangium; S, large spore within the sporangium wall w; s, the three aborted spores of the tetrad to which S belongs.

This type of fructification is the nearest approach to seed and pollen grains reached by any of the Pteridophytes, and its appearance at a time when the Lycopods were one of the dominant families is suggestive of the effect that such a position has on the families occupying it, however lowly they may be. The simple Pteridophyte Lycopods had not only the tall trunks and solid woody structure of a modern tree, but also a semblance of its seeds. Whether this line of development ever led on to any of the higher families is still uncertain. The feeling of most specialists is that it did not; but there are not wanting men who support the view that the lycopod affinity evolved in time and entered the ranks of the higher plants, and indeed there are many points of superficial likeness between the palæozoic Lycopods and the Coniferæ. Judged from their internal structure, however, the series through the ferns and Pteridosperms leads much more convincingly to the seed plants.

In their roots, or rather in the underground structures commonly called roots, the Lepidodendrons were also remarkable. Even more symmetrically than in their above-ground branching, the base of their trunks divided; there were four main large divisions, each of which branched into two and these into two again. These structures were called Stigmaria, and were common to all species of Lepidodendron and also the group of Sigillaria. On these horizontally running structures small appendages were borne all over their surface in great profusion, which were, both in their function and microscopic structure, rootlets. They left circular scars of a characteristic appearance on the big trunks, of which they were the only appendages. These scars show clearly on the fragments along the ledge to the left of the photograph. The exact morphological nature of the big axes is not known; their anatomy is not like that of roots, but is that of a stem, yet they do not bear what practically every stem, whether underground or not, has developed, namely leaves, or scales representing reduced leaves. Their nature has been commented on previously, and we cannot discuss the point further, but must be content to consider them as a form of root-bearing stem, practically confined to the Lycopods and principally developed among the palæozoic fossils of that group.

fig101

Fig. 101.:Transverse Section through a Rootlet of Stigmaria

oc, Outer cortex; s, space; ic, inner cortex; w, wood of vascular strand (wood only preserved); px, protoxylem group.

In microscopic structure the rootlets are extremely well known, because in their growth they have penetrated the masses of the tissues of other plants which were being petrified and have become petrified with them. The mass of decaying vegetable tissue on which the living plants of the period flourished were everywhere pierced by these intrusive rootlets, and they are found petrified inside otherwise perfect seeds, in the hearts of woody stems, in leaves and sporangia, and sometimes even inside each other!nFig. 95 shows such a root r lying in the space left by the decay of the soft tissue of the inner cortex in an otherwise excellently preserved Lepidodendron stem. In fig. 101 their simple structure is seen. They are often extremely irregular in shape, owing to the way they seem to have twisted and flattened themselves in order to fit into the tissues they were penetrating. No root hairs seem to have been developed in these rootlets, but otherwise their structure is that of a typical simple root, and very like the swamp-penetrating rootlets of the living Isoetes.

The Stigmarian axes and their rootlets are very commonly found in the “underclays” and “gannister” beds which lie below the coal seams, and they may sometimes be seen attached to a bit of the trunk growing upwards through the layers. They and the aerial stems of Lepidodendron are perhaps the commonest and most widely known of fossil plants.

Before leaving the palæozoic Lycopods another genus must be mentioned, which is also a widely spread and important one, though it is less well known than its contemporary. The genus Sigillaria is best known by its impressions and casts of stems covered by leaf scars. The stems were sometimes deeply ribbed, and the leaf scars were arranged in rows and were more or less hexagonal in outline, as is seen in fig. 102, which shows a cast and its reverse of the stem of a typical Sigillaria.

fig102

Fig. 102.:Cast and Reverse of Leaf Scars of Sigillaria. In A the shape of the leaf bases is clearly shown, the central markings in each being the scar of the vascular bundle and parichnos

In its primary wood Sigillaria differed from Lepidodendron in being more remote from the type with a primary solid stele. Its woody structure was that of a ring, in some cases irregularly broken up into crescent-shaped bundles. The secondary wood was quite similar to that of Lepidodendron.

Stigmaria and its rootlets belong equally to the two plants, and hitherto it has been impossible to tell whether any given specimen of Stigmaria had belonged to a Lepidodendron or a Sigillaria. Between the two genera there certainly existed the closest affinity and similarity in general appearance.

These two genera represent the climax of development of the Lycopod family. In the Lower Mesozoic some large forms are still found, but all through the Mesozoic periods the group dwindled, and in the Tertiary little is known of it, and it seems to have taken the retiring position it occupies to-day.