CHAPTER IX.

PTERIDOPHYTA (Vascular Cryptogams).

The Pteridophytes include plants which vary in size from a few millimetres[482] to several metres in height. The spore on germination gives rise to a small thalloid structure, the _prothallium_, on which the sexual organs are developed; this is the _gametophyte_ or sexual generation. The sexual organs have the form of typical archegonia and antheridia. From the fertilised egg-cell there is developed the Pteridophyte plant or _sporophyte_, which bears the spores. This asexual generation shows a well-marked external differentiation into stem and leaves, and bears true roots. Internally the tissues exhibit a high degree of differentiation into distinct tissue-systems. True vascular bundles occur, which may or may not be capable of secondary thickening by means of a _cambium_, _i.e._ a definitely localised zone of meristematic tissue. The sporangia are borne either on the ordinary foliage leaves or on special spore-bearing leaves called _sporophylls_, which differ in a greater or less degree from the sterile leaves.

The majority of the best known and most important Palaeozoic genera are either true Vascular Cryptogams, or possess certain of the pteridophytic characteristics combined with those of higher plants. It is not merely the commoner and more familiar recent genera with which the student of extinct types must be acquainted, but it is extremely important that he should make himself familiar with the rarer, less known and more isolated recent forms, which often throw most light on the affinities of the older representatives of the group. It is often the case, the more isolated living plants are, the more likely are they to afford valuable assistance in the interpretation of genera representing a class, which reached its maximum development in the earlier periods of the earth’s history. The importance of paying special attention to such recent plants as may be looked upon as survivals of a class now tending towards extinction, will be more thoroughly realised after the extinct vascular cryptogams have been dealt with.

A comparison of the Pteridophyta and Bryophyta brings out certain points of divergence. In the first place, the sporophyte assumes in the former class a much more prominent rôle, and the gametophyte has suffered very considerable reduction. The gametophyte, _i.e._ the structure which is formed on the germination of the asexually-produced spore, is usually short-lived, small, and more or less dependent on the sporophyte for its nutrition. In a few cases only is it capable of providing itself with the essential elements of food. On the other hand, the sporophyte, at a very early stage of its development becomes free from the gametophyte and is entirely self-supporting. Reproduction is effected as in the Bryophyta by sexual reproductive organs and by asexual methods. Not only have we in the Pteridophytes a much more complete external division of the plant-body into definite members, which subserve distinct functions, and behave as well-defined physiological organs adapted for taking a certain share in the life-functions of the individual, but the internal differentiation has reached a much higher stage. True vascular tissue, consisting of xylem and phloem, occurs for the first time in this class. The whole plant is traversed by one or more vascular strands composed of xylem and phloem elements, which are respectively concerned with the distribution of inorganic and organic food substances.

The Pteridophyta include the most important fossil plants. It is from a study of the internal structure of various extinct representatives of this class, that palaeobotanists have been able to contribute facts of the greatest interest and importance towards the advancement of botanical science.

The botanist’s chief aim in the anatomical investigation of Palaeozoic genera is to discover data which point the way to a solution of the problems of plant-evolution. In the abundant material afforded by the petrified remnants of ancient floras we have the means of tracing the past history of existing groups or individual forms, and it is from the Palaeozoic Pteridophytes that our most valuable results have been so far obtained.

In this and the following chapters of Volume I. two divisions of the Pteridophyta are dealt with in such detail as the nature of the book allows. In the earlier chapters of Volume II. the remaining representatives of this class will be described. As in the preceding chapters such recent plants will be described as are most essential for the correct interpretation of the fossil forms.

It is impossible to do more than confine our attention to a few only of the genera of living plants which directly concern us; some acquaintance with the general facts of plant morphology must be assumed. Among the most useful text-books or books of reference on the Pteridophyta the student may consult those mentioned in the footnote[483].

I. EQUISETALES.

Leaves usually small in proportion to the size of the whole plant, arranged in whorls at the nodes. Sporangia borne on specially modified sporophylls or sporangiophores, which are aggregated to form a definite strobilus or spore-bearing cone.

EQUISETACEAE. (Recent Species.)

The leaves are in whorls, coherent in the form of a sheath, and traversed by longitudinal veins which do not fork or anastomose. The stem is divided into comparatively long internodes separated by the leaf-bearing nodes, and the branches arise in the leaf-axils at the nodes. The fertile leaves or sporophylls differ from the sterile leaves, and usually occur in definite aggregations or strobili containing spores of one kind (_isosporous_). In the single living genus _Equisetum_, the outer coat of the mature spore forms two hygroscopically sensitive filamentous structures or _elaters_. On the germination of the spore the gametophyte is developed in the form of a small lobed prothallium 1–2 cm. in length. In most cases there are distinct male and female prothallia.

The genus _Equisetum_ L., the common Horse-tail, is the sole living representative of this Family. It occurs as a common native plant in Britain, and has a wide geographical distribution. Species of _Equisetum_ are abundant in the temperate zones of both hemispheres, and occur in arctic as well as tropical latitudes. Wallace[484] speaks of Horse-tails, “very like our own species,” growing at a height of 5000 feet on the Pangerango mountain in Java. In favourable situations the large British Horse-tail, _Equisetum maximum_ Lam. (= _E. Telmateia_ Erhb.), occasionally reaches a height of about six feet, and growing in thick clusters forms miniature forests of trees with slender erect stems and regular circles of long and thin branches. A tropical species, _Equisetum giganteum_ Linn.[2] living in the marshes of Mexico and Cuba[485], and extending southward to Buenos Ayres and Chili, reaches a height of twenty to forty feet, but the stem always remains slender, and does not exceed an inch in diameter. Groves of such tall slender plants on the eastern slopes of the Andes[486] suggest to the palaeobotanist an enfeebled forest-growth recalling the arborescent Calamites of a Palaeozoic vegetation. The twenty-five existing species of _Equisetum_ are remnants of various generic types of former epochs, and possess a special interest from the point of view of the geological history of plants. A brief description of the main characters of the recent genus will enable the student to appreciate the points of difference and agreement between the extinct and present representatives of the Equisetales.

[Illustration: FIG. 52. _Equisetum maximum_ Lam. A. Fertile shoot with strobilus and sterile leaf-sheaths [after Luerssen (89); slightly less than nat. size]. B. Sporophyll bearing open sporangia (after Luerssen; slightly enlarged). C. Part of a transverse section (diagrammatic); _v_, vallecular canals, _e_, endodermis, _c_, carinal canals (after Luerssen; × 20). D. _Equisetum arvense_ L. Part of a transverse section of an internode of a sterile shoot. _v_, cortex, _e_, endodermis, _x_, xylem tracheids, _a_ remains of annular tracheids of the protoxylem, _c_, carinal canal (after Strasburger; × 90).]

_Equisetum._

The plant consists of a perennial underground creeping rhizome, branching into secondary rhizomes, divided into well-marked nodes and internodes. From the nodes are given off two sets of buds, which may develope into ascending aerial shoots or descending roots. At each node is a leaf-sheath more or less deeply divided along the upper margin into teeth representing the tips of coherent leaves (fig. 52, A).

In some species one or more internodes of underground branches become considerably swollen and assume the form of ovate or elliptical starch-storing tubers, which are capable of giving rise to new plants by vegetative reproduction. Tubers, either singly or in chains, occur in _E. arvense_ Linn., _E. sylvaticum_ Linn., _E. maximum_ Lam., among British species.

[Illustration: FIG. 53. Rhizome (R) of _Equisetum palustre_ L. with a thin shoot giving off roots and tuberous branches from a node [after Duval-Jouve (64)].]

In the example shown in fig. 53 (_Equisetum palustre_ L.[487]) the stout rhizome R gives off from its node, marked by a small and irregular leaf-sheath, two thin roots and a single shoot. The latter has a leaf-sheath at its base, and from the second node, with a larger leaf-sheath, there have been developed branches with tuberous internodes; the constrictions between the tubers and the tips of the terminal tubers bear small leaf-sheaths. Branched roots are also given off from the upper node of the erect shoot.

Near the surface of the ground the buds on the rhizome nodes develope into green erect shoots. The shoot axis is marked out into long internodes separated by nodes bearing the leaf-sheaths. The surface of each internode is traversed by regular and more or less prominent longitudinal ridges and grooves; each ridge marking the position of an internal longitudinal vascular strand. In the axil of each leaf, that is in the axil of each portion of a leaf-sheath corresponding to a marginal uni-nerved tooth, there is produced a lateral bud which may either remain dormant or break through the leaf-sheath and emerge as a lateral branch. At the base of each branch an adventitious root may be formed from a cell immediately below the first leaf-sheath, but in aerial shoots the roots usually remain undeveloped. The lateral branches repeat on a smaller scale the general features of the main axis. In some species, the shoots are unbranched, and in others the slender branches arise in crowded whorls from each node. Leaves, roots and branches are given off in whorls, and the whorls from each node alternate with those from the node next above and next below.

[Sidenote: ANATOMY OF EQUISETUM.]

In some species of _Equisetum_ the aerial stem terminates in a conical group of sporophylls, while in others the strobilus is formed at the apex of a pale-coloured fertile shoot, which never attains any considerable length and dies down early in the season of growth (fig. 52, A). Below the terminal cone or strobilus there occur one or two modified leaf-sheaths. Such a ring of incompletely developed leaves intervening between the cone of sporangiophores and the normal leaves, is known as the _annulus_. The annulus is seen in fig. 52, A, immediately below the lowest whorl of sporophylls; it has the form of a low sheath with a ragged margin. In the region of the cone the internodes remain shorter, and the whorls of appendages, known as sporophylls or sporangiophores, have the form of stalked stru