FOSSIL PLANTS.

CAMBRIDGE UNIVERSITY PRESS
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C. F. CLAY, Manager

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Part of a transverse section of a Permian Osmundaceous Fern stem, Thamnopteris Schlechtendalii (Eichwald). a, outer xylem; b, inner xylem. For description, see [page 329]. (After Kidston and Gwynne-Vaughan. Very slightly reduced.)

FOSSIL PLANTS

A TEXT-BOOK FOR STUDENTS
OF BOTANY AND GEOLOGY

BY

A. C. SEWARD, M.A., F.RS.
PROFESSOR OF BOTANY IN THE UNIVERSITY; FELLOW OF ST JOHN’S
COLLEGE AND HONORARY FELLOW OF EMMANUEL COLLEGE, CAMBRIDGE

WITH 265 ILLUSTRATIONS

VOL. II

CAMBRIDGE:
AT THE UNIVERSITY PRESS
1910

Cambridge:
PRINTED BY JOHN CLAY, M.A.
AT THE UNIVERSITY PRESS.

PREFACE

I REGRET that pressure of other work has prevented the completion of this Volume within a reasonable time since the publication of Volume I. Had Volume II been written ten years ago, the discoveries made in the course of the last decade would have given an out-of-date character to much of the subject-matter. It is more especially in regard to the Ferns and the extinct members of the Gymnosperms that our outlook has been materially altered by recent contributions to Palaeobotany. It is, however, some satisfaction to be able to add that recent progress has been relatively slight in that part of the subject dealt with in the first volume.

The original intention was to complete the whole work in two volumes. Soon after the second volume was begun, it became evident that the remaining divisions of the plant-kingdom could not be included within the compass of a single volume. I decided, therefore, to take the consequences of having embarked on too ambitious a plan of treatment, and to preserve uniformity of proportion by reserving the seed-bearing plants for a third volume. The third volume will include the Pteridosperms, other than those briefly described in the final chapter of the present volume, and other classes of Gymnosperms. I propose also to devote such space as is available within the limits of a text-book to the neglected subject of the geographical distribution of plants at different stages in the history of the earth. It is my intention to complete Volume III with as little delay as possible. As I have written elsewhere, the past history of the Flowering plants needs special treatment, and anything more than a mere compilation can be adequately attempted only after considerable research and with the assistance of botanists possessing a special knowledge of different families of Angiosperms. The need of a critical examination of available data in regard to the geological history of this dominant group will not be lost sight of.

I am well aware that while certain genera have received an undue share of attention in the present volume, others have been ignored or treated with scant consideration. For this inconsistency I have no excuse to offer, beyond the statement that the subject is a large one, and selection is necessary even though the work consists of three volumes.

The publication in 1909 of a collection of excellent photographs of Palaeozoic Plants, with brief descriptive notes, by Mr Newell Arber, as one of a series of popular “Nature Books,” bears striking testimony to the remarkable spread of interest in the study of the vegetation of the past, which is one of the outstanding features in the recent history of botanical science.

In the list of illustrations I have mentioned the source of all figures which have been previously published. I would, however, supplement the statement of fact with an expression of thanks to corporate bodies and to individuals who have allowed me to make use of blocks, drawings, or photographs.

I wish to thank my colleague, Mr A. G. Tansley, for placing at my disposal several blocks originally published in the pages of the New Phytologist. To Professor Bertrand of Lille and to his son Dr Paul Bertrand I am indebted for several prints and descriptive notes of specimens in their possession. My friends Dr Nathorst of Stockholm and Dr Zeiller of Paris have generously responded to my requests for information on various points. I wish especially to thank Dr Kidston for several excellent prints of specimens in his collection and for the loan of sections. I have profited by more than one examination of his splendid collection at Stirling. Professor Weiss has generously allowed me to borrow sections from the Manchester University collections, more especially several which have been reproduced in the chapter devoted to the genus Lepidodendron. To Professor F. W. Oliver my thanks are due for the loan of sections from the collection under his charge at University College. I have pleasure also in thanking Dr Scott, not only for lending me sections of a Lepidodendron and for allowing me to use some drawings of Miadesmia originally made by Mrs Scott for reproduction in his invaluable book, Studies in Fossil Botany, but for kindly undertaking the laborious task of reading the proofs of this volume. It would be unfair to express my gratitude to Dr Scott for many helpful suggestions and criticisms, without explicitly stating that thanks to a friend for reading proofs must not be interpreted as an attempt to claim his support for all statements or views expressed. The General Editor of the Series, Mr A. E. Shipley, has also kindly read the proofs. I am under obligations also for assistance of various kinds to Prof. Thomas of Auckland, New Zealand, to Mr Boodle of Kew, to Mr D. M. S. Watson of Manchester, to Mr T. G. Hill of University College, and to Mr Gordon of Emmanuel College, Cambridge. I am indebted to the kind offices of Miss M. C. Knowles for the photograph of the specimen of Archaeopteris hibernica in the Irish National Museum, Dublin, reproduced on [page 561].

Many of the illustrations are reproduced from drawings by my wife: those made from the actual specimens are distinguished by the addition of the initials M. S. I am grateful to her also for some improvements in the letter-press. For the drawings made from sections and for some of the outline sketches I am responsible. I have availed myself freely of the facilities afforded by Professor McKenny Hughes in the Sedgwick Museum of Geology for the examination of specimens under the charge of Mr Newell Arber, the University Demonstrator in Palaeobotany. It is a pleasure to add that, as on former occasions, I am indebted to the vigilance of the Readers of the University Press for the detection of several errors which escaped my notice in the revision of the proofs.

A. C. SEWARD.

Botany School, Cambridge.
March 12, 1910.

ERRATA IN VOL. I

• Page 16, line 4. For “The North American Tulip tree” read The Tulip tree of North America and China.
• „ 66, line 2 from the bottom. For “Browera” read Berowra.
• „ 127, line 3 and 4 from bottom. For Achyla and Palaeachyla read Achlya and Palaeachlya.
• „ 145, lines 4 and 5. For “Upper Greensand” read Lower Eocene.
• „ 162, line 3 from bottom. For “Corallina barbata” read Cymopolia barbata.
• „ 170, line 20. For “sporangiaphore” read sporangiophore.
• „ 185, line 2. The genera Udotea and Halimeda, members of the Siphoneae, are incorrectly included under the Corallinaceae.
• „ 191, line 11 from bottom. Omit Chondrus crispus, which is one of the Florideae and not a Brown Alga.
• „ 202, line 13. For “Halmeda” read Halimeda.
• „ 250, line 11. For “three” read the.
• „ 381, line 10. For “Calamopytus” read Calamopitys.

CHAPTER XII[1].

SPHENOPHYLLALES (concluded).

[Sphenophyllum].

The account of the Sphenophyllales given in the first volume[2] of this work must be extended and somewhat modified in the light of recent work on the fertile shoots of Sphenophyllum.

Sphenophyllostachys Dawsoni (Will.) was described as consisting of an axis bearing superposed whorls of bracts connate at the base in the form of a shallow funnel-shaped collar giving off from the upper surface and close to the axis of the cone two concentric series of sporangiophores. Occasionally there are three series, as represented in [fig. 112]. In another type of strobilus, Sphenophyllostachys Römeri[3] each sporangiophore terminates in two pendulous sporangia ([fig. 113], A; see also fig. 107, C, vol. I.). It has already been pointed out that the common occurrence of detached strobili necessitates their description under distinct specific names; it is only by a rare accident that we can assign fossil cones to their vegetative shoots. There are, however, reasons for believing that Sphenophyllostachys Dawsoni is the strobilus of the plant originally described by Sternberg[4] from impressions of foliage-shoots as Rotularia cuneifolia. Another difficulty presented by petrified material is that of determining, with certainty, whether two imperfect specimens, differing from one another in features which do not appear to be of sufficient importance to warrant specific separation, are forms of one species or portions of specifically distinct cones. It has been pointed out by Scott[5] that the strobilus known as Sphenophyllostachys Dawsoni probably includes two distinct species, one being the cone of Sphenophyllum cuneifolium Sternb., and the other the cone of S. myriophyllum Crép[6]. The stem of S. myriophyllum agrees anatomically with the type known as Sphenophyllum plurifoliatum Will. and Scott[7].

Fig. 112. Sketch of a radial longitudinal section of Sphenophyllostachys. There are usually two concentric series of sporangia on the sporophylls, not three as shown in the figure. The upper figure (after Zeiller) shows the linear bracts in surface-view.

In addition to the two types of cone already mentioned, Sphenophyllostachys Dawsoni and S. Römeri, others have been described by Kidston from carbonised impressions. One of these is the fertile branch of Sphenophyllum majus[8]. The basal portions of the bracts of each whorl form a narrow collar round the axis of the cone; the free portion of each bract consists of a lamina divided into two equal bifid lobes bearing on its upper surface one group, or possibly two groups, of four sessile sporangia between the narrow coherent bases of the laminae and the sinus between the terminal lobes ([fig. 113], C). Another characteristic feature is the greater length of the internodes; this renders the cone less compact and less sharply differentiated from the vegetative shoots than those of other species. A specimen in Dr Kidston’s collection illustrates the peculiar character of the fertile portion of this species; it consists of an axis bearing a succession of lax sporophylls succeeded above and below by whorls of sterile leaves. In this species, therefore, we cannot speak of a compact strobilus at the end of a shoot of limited growth, but of axes in which sterile and fertile leaves are borne alternately[9], a condition recalling the alternation of foliage leaves and sporophylls in Tmesipteris and in Lycopodium Selago.

Fig. 113.

1. Sphenophyllostachys Römeri. (Solms-Laubach.)
2. Sphenophyllum trichomatosum Stur.
3. Sphenophyllum majus. Bronn. (A–C. After Kidston.)

Another form of cone, also from the Middle Coal Measures, is referred by Kidston to Sphenophyllum trichomatosum Stur[10] ([fig. 113], B): this is characterised by the more horizontal position of the bracts, which “do not appear to be so much or so suddenly bent upwards in their distal portion as in some other species of Sphenophyllum,” and by sessile sporangia borne singly on the upper face of each bract.

Fig. 114. Sphenophyllostachys fertilis (Scott). (After Scott.) Diagram of a node in longitudinal section, showing one sporophyll and the base of the opposite one. v.l. ventral lobe of sporophyll; v.s. one of the segments into which it divides; v.s′. stump of another segment; d.l. dorsal lobe; d.s., d.s′. segments of dorsal lobe.

A more recent addition to our knowledge of the fertile shoots of Sphenophyllum is due to Scott who has described a new type of cone under the name Sphenophyllum fertile[11]. The petrified specimen on which the species was founded was discovered by Mr James Lomax in the Lower Coal Measures of Lancashire; it represents a portion of a cone 6 cm. long and approximately 12 mm. broad. The axis contains a single vascular cylinder agreeing in essentials with the type of stem structure known as Sphenophyllum plurifoliatum. The nodal regions, which exhibit the slight swelling characteristic of the genus, bear several (probably twelve) appendages connate at the base and forming a narrow flange encircling the axis. Each bract, the base of which forms part of the narrow collar surrounding the axis, consists of two lobes, ventral and dorsal, divided palmately into several (sometimes four) segments or sporangiophores ([fig. 115]). Each sporangiophore terminates distally in an oblong or oval lamina bearing two sporangia on its adaxial face ([fig. 114]). The space between the axis and the periphery of the cone is thus occupied by crowded peltate laminae, each with its pair of sporangia. A single vascular bundle supplies each sporangiophore and bifurcates in the distal lamina into two branches which extend to the bases of the sporangia. The sporangia agree in structure with those of other species of Sphenophyllum: the spores are of one size and elliptical, characterised by the presence of several sharp ridges or flanges encircling the spore-wall in the direction of the major-axis. Sphenophyllostachys fertilis differs from all previously recorded types in the absence of sterile bracts. The appendages of the cone-axis are all fertile, a striking contrast to the differentiation into protective and sporangia-bearing bracts which constitutes a constant feature in the cones of Sphenophyllum and Calamites. It is possible, as Scott suggests, that the absence of sterile segments is the result of modification of the more usual type of strobilus; instead of the dorsal and ventral lobes of the bracts sharing between them the duties of protection and spore-production, the whole of each bract is constructed on the plan of the maximum spore-output, the laminar terminations of the sporangiophores serving the purpose of protection. The cone may be described as more specialised than the normal type of strobilus for reproductive purposes[12].

Fig. 115. Sphenophyllostachys fertilis (Scott). (After Scott.) Diagram of a single sporophyll as it would appear in a transverse section of the cone; showing one lobe (dorsal or ventral). ax, part of axis to which the sporophylls are attached.

Fig. 116. Sphenophyllostachys Dawsoni. (After Thoday.) A. Larger spores; B, abortive spores; C, mature spores showing the characteristic spines.

It has been stated, on evidence which is unsatisfactory, that Sphenophyllum possesses two kinds of spores. While regarding the genus as homosporous on the evidence before us, it is interesting to find that cases occur in which the spores in the same sporangium exhibit a marked difference in size. Attention has been called by Williamson and Scott[13] to variation in the dimensions of spores: a more pronounced difference in size has been recorded by Mr Thoday[14] who gives 120μ as the maximum and 90μ as the minimum diameter of the spores in a cone of Sphenophyllostachys Dawsoni. The presence of several abortive spores in the sporangium ([fig. 116]) containing the larger spores favours the view that this difference in size may be the first step towards the development of heterospory.

It is clear that the types of strobilus designated Sphenophyllostachys ([figs. 112–114]) present a divergence of characters too great to be comprised under one genus; but in the absence of fuller information, we cannot do otherwise than follow the only logical custom of grouping them together as examples of strobili borne by plants which, in the present state of our knowledge, are most conveniently referred to the genus Sphenophyllum.

Cheirostrobus.

This generic name was applied by Dr Scott[15] to a calcified cone obtained by Mr James Bennie in 1883 from the Lower Carboniferous plant-beds of Pettycur near Burntisland on the Firth of Forth. Cheirostrobus is distinguished from Sphenophyllostachys by its greater breadth (3.5 cm.); externally it agrees more closely with the fertile shoots of Lepidodendron than with those of Sphenophyllum. A single vascular cylinder having the form of a fluted Doric column ([fig. 117], B, x) occupies the axis of the cone: it consists for the most part of reticulate tracheae which tend to assume a short or isodiametric form in the central region; the smaller protoxylem tracheids with the spiral form of pitting constitute the sharp and prominent ridges at the periphery of the xylem-cylinder. In the outer part of the cylinder the metaxylem[16] consists exclusively of tracheae, but towards the centre of the axis these are associated with numerous parenchymatous cells.

The xylem is therefore centripetal in origin as in Sphenophyllum and in nearly all recent and fossil members of the Lycopodiales. In the type-specimen of Cheirostrobus the vascular cylinder of the cone consists entirely of primary xylem, but secondary xylem has been found in a more recently discovered specimen[17]. Secondary xylem occurs also in the peduncle of the cone. No appreciable remains of phloem have been found. The cortex consists of slightly elongated rather thick-walled tissue containing secretory sacs. Crowded superposed whorls of bracts (or sporophylls), usually twelve in each whorl, are borne on the axis and each sporophyll receives a single vascular bundle from one of the vertical ridges of the xylem column ([fig. 117], A, lt). The members of each whorl are connate at the base: from this narrow collar each sporophyll branches into an upper or dorsal and a lower or ventral limb ([fig. 117], A, f and s). Each limb divides palmately at a short distance from its origin into three slender segments, which extend in a horizontal direction and terminate in large laminar expansions ([fig. 117], B, s) to afford a protective covering to the surface of the cone. The upper set of three segments, constituting sporangiophores ([fig. 117], A, B, f) or fertile divisions of the sporophyll, expand distally into comparatively bulky laminae; each of these bears on its adaxial face four diagonally placed outgrowths which form the short pedicels of very long and narrow sporangia. The three lower segments—the sterile divisions of the sporophylls—([fig. 117], A, B, s) are similar to the upper set except in their greater length and in the kite-shaped form of their distal laminae which are provided with lateral lobes. The single vascular strand which supplies each sporophyll is represented at lt in [fig. 117], B; at lt′ the strand has divided into four, the three upper bundles in the figure supply the sterile segments and the single lower bundle ultimately divides into three which supply the fertile segments. A pair of blunt processes (fig. A, s) extend downwards over the ends of the underlying fertile lamina and two slender prolongations extend upwards through several internodes.

Fig. 117. A, B. Cheirostrobus pettycurensis Scott. (After Scott.)
C, D. Pseudobornia ursina Nath. (After Nathorst.)

1. Diagrammatic radial longitudinal section of part of the cone-axis and two sporophylls. lt, bundle passing out to sporophyll; f, fertile segment of sporophyll showing two sporangia; s, sterile (lower) segment.
2. Part of transverse section. x, stele; lt, lt′, bundles on their way to sporophylls; a, tips of sterile segments of lower sporophylls.
3. Palmately branched leaf (½ natural size).
4. Node of stem showing leaf-bases.

An economical arrangement of the long and narrow sporangia and of the sporophyll-segments between the axis and the periphery of the cone is rendered possible by the interlocking of the sterile and fertile segments by means of a groove in the upper face of the latter for the accommodation of the former. The sporangia are characterised by their unusually long and narrow form: the length of a sporangium may reach 1 centimetre. In the structure of the wall the sporangia of Cheirostrobus agree closely with those of Calamostachys[18] and Sphenophyllostachys. The spores are of one size only. The vascular cylinder of the peduncle, originally described by Williamson[19] as the peduncle of a large Lepidostrobus (the cone of Lepidodendron), is characterised by the presence of a short radially disposed zone of secondary tracheids, a feature, as Scott points out, which may extend into the axis of the cone. It is noteworthy that the protoxylem elements are not always external, but occasionally occur internal to one or two of the outermost metaxylem tracheae: the usual exarch[20] structure of the central cylinder is not therefore absolutely constant, but may be replaced by a mesarch arrangement.

The presence of a few sterile leaves on the peduncle below the fertile portion of the cone, which agree in their lobed laminae with the sporophylls, is the only fact which we possess as to the form of the vegetative characters of the genus.

The above description is sufficient to indicate the extraordinary complexity and high degree of specialisation of Cheirostrobus. The sporophylls, with their trilobed segments, and the crowded sporangia of exceptional length attached only by a narrow base constitute striking peculiarities of the genus.

It is unfortunate that we are still without any satisfactory evidence as to the nature of the plant the cones of which have been made the type of a new genus and a new family. Cheirostrobus affords an interesting example of a type of reproductive shoot constructed on a plan sui generis, and may be classed with some other extinct genera as instances of the production in the course of evolution of architectural schemes which appear to have been ill adapted for competition with equally efficient though much simpler types. But the discovery of these isolated forms of restricted geological range among the relics of the Palaeozoic vegetation frequently supplies a key to phylogenetic problems. Cheirostrobus by its complex combination of features characteristic of the Equisetales, the Lycopodiales and the genus Sphenophyllum throws a welcome light on the inter-relationships of groups which represent divergent series. The combination of morphological features in this generalised type led the author of the genus to describe it as a descendant of an old stock which existed prior to the divergence of the Equisetales and Lycopodiales.

The discovery of this new type of strobilus naturally led to a search among Lower Carboniferous plants for vegetative shoots exhibiting characters conformable with the whorled and branched leaves of Cheirostrobus. In Sphenophyllum we have a genus obviously comparable with Cheirostrobus as regards the form and disposition of the leaves, but the differences between the cones and the striking similarity of the vascular cylinder of the latter to that of Lepidodendron demonstrate conclusively that we must look elsewhere for the vegetative members of the plant which produced cones of the Cheirostrobus type.

PSEUDOBORNIA

In 1902 Professor Nathorst[21] instituted the generic name Pseudobornia for plants of which imperfect examples had previously been referred by Heer[22] to Calamites under the name C. radiatus. Heer’s plants were obtained from Upper Devonian rocks of Bear Island in the Arctic seas and additional specimens were brought from the same locality by the Swedish Polar Expedition of 1898. Pseudobornia possesses jointed stems ([fig. 117], D) bearing whorled and shortly stalked leaves, often four in number, at each node. The leaves are palmately branched with fine serrated edges ([fig. 117], C). Certain specimens, which are no doubt correctly described by Nathorst as cones, are characterised by a thick axis bearing whorled leaves with sporangia on their lower surfaces, but the material is not sufficiently well preserved to render possible a recognition of structural details. It has been suggested by Scott that Pseudobornia may possibly be referable to the Sphenophyllales and that the stem of Cheirostrobus “may have had something in common with” Nathorst’s genus[23]. The beds in which the stems occur are of Upper Devonian age, while Cheirostrobus was found in Lower Carboniferous rocks: this difference in age is not, however, a serious objection to the validity of the comparison. We cannot do more than express the view that Pseudobornia, so far as can be ascertained without an examination of petrified material or of more perfect impressions of strobili, exhibits vegetative features not inconsistent with the morphological characters of the fertile shoots known as Cheirostrobus.

•••••

The institution of a special group-name for the reception of Sphenophyllum is justified by the sum of its morphological features, which do not sufficiently conform to those of any existing group of Pteridophytes to warrant its inclusion in a system of classification based on recent genera. In the case of Cheirostrobus we are limited to the characters of the cone and its peduncle. The suggestion that the Devonian fossils known as Pseudobornia may represent the foliage shoots of a plant closely related to Cheirostrobus has still to be proved correct. Although we may find justification in the highly complex and peculiar structure of Cheirostrobus for the recognition of the genus as a type of still another group of Pteridophytes, it would be unwise to take this step without additional knowledge.

The undoubted similarity between Cheirostrobus and Sphenophyllum coupled with striking points of difference favours the inclusion of the two genera in distinct families placed, for the present at least, in the group Sphenophyllales.

Group SPHENOPHYLLALES.

• Sphenophylleae: genus Sphenophyllum.
• Cheirostrobeae: genus Cheirostrobus.

It has recently been proposed to include the family Psilotaceae, comprising the two recent genera Psilotum and Tmesipteris, as another subdivision of the Sphenophyllales. This proposal had been made by Professor Thomas[24] primarily on the ground that the sporophylls of Tmesipteris and Psilotum appear to afford the closest parallel among existing plants to the peculiar form of sporophyll characteristic of the Sphenophyllales. The morphological interpretation of the sporophylls of both Sphenophyllum and Cheirostrobus has been the source of considerable discussion[25]. If we regard each sporophyll as a leaf with two lobes, one fertile and one sterile, except in the case of Sphenophyllostachys fertilis in which both are fertile, an obvious comparison may be made with the fern Ophioglossum; but the difference between a single fern frond, consisting of a comparatively large sterile lamina bearing a fertile branch composed of a long axis with two rows of sporangia embedded in its tissues, and the whorled sporophylls of Sphenophyllum is considerable.

PSILOTACEAE

A brief reference may be made to the principal reasons which have led to the suggestion that the Psilotaceae should be included in the Sphenophyllales. The shoots of Tmesipteris bear simple foliage leaves spirally disposed on a slender axis, and in association with these occur sporophylls consisting of a short axis bearing a pair of small lobes and a bilocular synangium[26] ([fig. 120], B). The synangium is seated on a very short stalk given off from its sporophyll at the base of the pair of laminae: the synangium with its short stalk may be spoken of as the sporangiophore. In most cases the synangium appears to be sessile on the sporophyll, but occasionally the much reduced stalk is prolonged and forms an obvious feature. Dr Scott[27] suggested that the Tmesipteris synangium with its axis may correspond to the ventral lobe (or sporangiophore) of Sphenophyllum. In the latter genus the whorled sporophylls consist in most species of a dorsal and a ventral lobe, the latter serving as a sporangiophore bearing one or more sporangia; in Tmesipteris the sporophylls are spirally disposed and each consists of a bilobed sterile portion bearing a septate sporangium or bilocular synangium on a very short ventral lobe. Professor Bower[28], in his account of the development and structure of the sporophylls of Tmesipteris, drew attention to the comparatively frequent occurrence of abnormal sporophylls and spoke of the plant as unstable. More recently Professor Thomas[29] of Auckland has carefully examined living plants, with the result that variations of different kinds are proved to be exceedingly common. He finds that sporophylls occur which exhibit repeated dichotomy of the axis ([fig. 120], D, F) and thus each may bear four instead of two leaf-lobes and three synangia, one at the first fork and one at each of the forks of the second order[30].

Other abnormalities occur in which the synangium is raised on a distinct stalk instead of being more or less sessile at the point from which the leaf-lobes diverge. A third form of departure from the normal is that in which there is no synangium on the bilobed sporophyll, its place being taken by a leaf-lobe. The deduction from the occurrence of these abnormalities is that the synangium of Tmesipteris represents a ventral leaf-lobe, as Scott suggested. Professor Thomas draws attention to the resemblance between Tmesipteris sporophylls and the foliage-leaves of Sphenophyllum, which are either simple with dichotomously branched veins or the lamina is deeply divided into two or more segments. In some types of Sphenophyllostachys the bracts are simple (S. Dawsoni), but in others (Sphenophyllum majus, [fig. 113], C) they are forked like the foliage-leaves and bear a close resemblance to the abnormal sporophylls of Tmesipteris. Moreover, in Sphenophyllostachys Römeri ([fig. 113], A) each ventral lobe of a sporophyll bears two sporangia, a condition almost identical with that represented by the occasional occurrence of a synangium on a comparatively long stalk in Tmesipteris. Similarly the more elaborate sporophylls of Cheirostrobus may be compared with the branched sporophylls of Tmesipteris ([fig. 120]). This agreement between the sporophylls of the Palaeozoic and recent genera acquires additional importance from the very close resemblance between the exarch stele of Sphenophyllum and that of the genus Psilotum, which conforms to the Palaeozoic type not only in the centripetal character of the primary xylem and in its exarch structure, but also in the occasional occurrence of secondary xylem[31], and in the stellate form of its transverse section. The occasional mesarch structure of the stele of Cheirostrobus finds a parallel in the mesarch xylem groups in the stem of Tmesipteris. It is thus on the strength of these resemblances that Thomas and Bower would remove the Psilotaceae from the group Lycopodiales and unite them with Sphenophyllum and Cheirostrobus in the Sphenophyllales. While admitting the validity of the comparison briefly referred to above, I prefer to retain the Psilotaceae as a division of the Pteridophyta including only Psilotum and Tmesipteris.

SPHENOPHYLLUM

In his recent book on The Origin of Land Flora, Prof. Bower raises objection to the use of the term ventral lobe in speaking of the sporangium-bearing stalk or sporangiophore borne on the sporophyll of Sphenophyllum. He points out that the use of this term implies the derivation of the sporangiophore by metamorphosis of part of a vegetative leaf, an opinion untenable in the absence of proof. The designation sporangiophore is no doubt preferable to that of ventral lobe as it carries with it no admission of particular morphological value; as a further concession to a non-committal attitude we may provisionally at least regard a sporangiophore as an organ sui generis “and not the result of modification of any other part[32].”

The view put forward by Prof. Lignier[33] that the Sphenophyllales are descendants of primitive ferns is not convincing, and his comparison of Sphenophyllum with Archaeopteris lacks force in view of our ignorance as to the nature of the reproductive organs of the latter genus. That the Sphenophyllales are connected with the Equisetales and with the Psilotales by important morphological features is clear; but the comparison between the sporophylls of the extinct genera with those of the existing genus Tmesipteris, though helpful and possibly based on true homology, cannot be considered as settling the morphological value of the sporangiophores of Sphenophyllum and Cheirostrobus.

I do not propose to discuss at length the different views in regard to the morphological nature of the sporangiophore of Sphenophyllum. The comparison, which we owe in the first instance to Scott, with the synangium of the Psilotales with its short stalk, though not accepted by Lignier as a comparison based on true homology, is one which appeals to many botanists and is probably the best so far suggested. The further question, whether these sporangiophores are to be called foliar or axial structures is one which has been answered by several authors, but it is improbable that we shall soon arrive at a decision likely to be accepted as final. Discussions of this kind tend to assume an exaggerated importance and frequently carry with them the implication that every appendage of the nature of a sporangiophore can be labelled either shoot or leaf. We treat the question from an academic standpoint and run a risk of ignoring the fact that the conception of stem and leaf is based on morphological characteristics, which have been evolved as the result of gradual differentiation of parts of one originally homogeneous whole. There is much that is attractive in the view recently propounded by Mr Tansley that a leaf is not an appendicular organ differing ab initio from the axis on which it is borne, but that it is in phylogenetic origin a “branch-system of a primitive undifferentiated sporangium-bearing thallus[34].” Admitting the probability that this view is correct, our faith in the importance of discussions on the morphological nature of sporangiophores is shaken, and we realise the possibility that our zeal for formality and classification may lead to results inconsistent with an evolutionary standpoint[35].

CHAPTER XIII.

PSILOTALES.

The two recent genera Psilotum and Tmesipteris are usually spoken of as members of the family Psilotaceae which is included as one of the subdivisions of the Lycopodiales. It is probable, as Scott[36] first suggested, that these two plants are more nearly allied than are any other existing types to the Palaeozoic genus Sphenophyllum.

We may give expression to the undoubted resemblances between Tmesipteris and Psilotum and the Sphenophyllales by including the recent genera as members of that group, originally founded on the extinct genus Sphenophyllum; this is the course adopted by Thomas[37] and by Bower[38]: or we may emphasise the fact that these two recent genera differ in certain important respects from Lycopodium and Selaginella by removing them to a separate group, the Psilotales. The latter course is preferred on the ground that the inclusion of Psilotum and Tmesipteris in a group founded on an extinct and necessarily imperfectly known type, is based on insufficient evidence and carries with it an assumption of closer relationship than has been satisfactorily established.

The genus Tmesipteris ([fig. 120], A) is represented by a single species T. tannensis Bertr.[39] which usually occurs as an epiphyte on the stems of tree-ferns in Australia, New Zealand, and Polynesia. Psilotum, with two species P. triquetrum Sw. ([fig. 118]) and P. complanatum Sw., flourishes in moist tropical regions of both hemispheres, growing either on soil rich in organic substances or as an epiphyte. Both genera are considered to be more or less saprophytic.

Fig. 118. Psilotum triquetrum (½ natural size).

1. Synangium.
2. Sporophyll after removal of the synangium. (M.S.)

Psilotum. The common tropical species P. triquetrum ([fig. 118]) is characterised by an underground rhizome which forms a confused mass of dark brown branches covered with filamentous hairs as substitutes for roots and gives off erect repeatedly forked aerial shoots. In P. complanatum[40] the habit is similar to that of the more abundant and better-known species, but the pendulous shoots are characterised by their broader and flatter form. In both species the function of carbon-assimilation is performed by the outer cortex of the green branches, as the small size of the widely-separated foliage leaves renders them practically useless as assimilating organs.

The sporophylls consist of a short axis terminating in two small divergent forks and bearing on its adaxial surface a trilocular or in rare cases a bilocular synangium ([fig. 118], A and B). The walls of the loculi are composed of several layers of cells and dehiscence takes place along three lines radiating from the centre of the synangium. Professor Thomas[41] has recorded “fairly numerous instances in Psilotum of a second dichotomy of one branch of the first fork, or, less frequently, of both branches”: instead of one synangium subtended by the two slender leaflets of the forked sporophyll-axis, there may be two synangia and three leaf-lobes or three synangia and four leaf-lobes. The occurrence of both these abnormalities in Psilotum and Tmesipteris shows a decided tendency in the Psilotales to a repeated dichotomy of the sporophylls[42].

A single stele[43] with a fluted surface occupies the axis of an aerial shoot ([fig. 119], A); the axial region is occupied by a core of elongated mechanical elements (s), which may occasionally extend to the periphery of the xylem and break the continuity of the band of scalariform tracheae ([fig. 119], A, a). The tracheae form the arms of an irregularly stellate stele and each arm is terminated by protoxylem elements ([fig. 119], B, px). The rays of the xylem cylinder, which may be as many as six or eight in the upper part of the aerial shoots, become reduced in number as the rhizome is approached, assuming a diarch structure near the junction. In the rhizome the xylem forms an approximately triangular group of tracheae without any core of mechanical elements. Three to four layers of parenchyma succeeded externally by an ill-defined phloem ([fig. 119], A, p) surround the xylem and a fairly distinct endodermis ([fig. 119], A and B, e) encloses the whole. To Mr Boodle[44] is due the interesting discovery that in some parts of the rhizome the parenchymatous zone surrounding the scalariform tracheae may become the seat of meristematic activity which results in the production of secondary tracheae often characterised by a sinuous longitudinal course. There is no definite cambium, but the radially disposed tracheae and the adjacent parenchymatous elements clearly demonstrate the secondary nature of the tissue immediately external to the group of primary xylem. Fig. 119, C, drawn from a section kindly supplied by Mr Boodle, shows the secondary xylem elements at x² associated with radially disposed thin-walled cells abutting on the primary xylem, x¹. It is probable that this added tissue may be a remnant of a more extensive secondary thickening characteristic of the ancestors of the recent species. In their manner of occurrence and sinuous course these secondary tracheids bear a resemblance to the secondary xylem of Lepidodendron fuliginosum[45]. The stele of the aerial shoot bears a fairly close resemblance to the vascular axis of Cheirostrobus, and its three-rayed form in the lower portions of the green branches recalls that of the Sphenophyllum stele, except that the axial xylem elements of the Palaeozoic genus are usually represented in Psilotum by mechanical tissue. The cortex consists of three regions ([fig. 119], A), an outer zone of chlorophyllous tissue (a) rich in intercellular spaces succeeded by a band of mechanical tissue (b) which gradually passes into an inner region of larger and thinner-walled cells (c).

Fig. 119.

1. Diagram of transverse section of aerial shoot of Psilotum triquetrum. a—c, cortex; p, phloem; e, endodermis; s, stereome; x, xylem; a, gap in xylem.
2. Enlarged view of one of the angles of the xylem shown in A. px, protoxylem.
3. Part of transverse section of an approximately triangular rhizome stele showing a portion of the metaxylem x¹; px, protoxylem elements; x², secondary xylem.

TMESIPTERIS

The genus Tmesipteris[46] agrees with Psilotum in general habit and in its epiphytic and probably in some degree saprophytic mode of life. Its brown rootless rhizome, which grows among the roots of tree-ferns or rarely in the ground, gives off pendulous or erect shoots reaching a length of two feet and bearing lanceolate mucronate leaves 2–3 cm. long ([fig. 120], A) attached by decurrent leaf-bases. The sporophylls, replacing the upper leaves or occurring in more or less well-defined zones alternating with the foliage leaves, consist of a short axis terminating in a pair of lanceolate lobes and bearing on its adaxial surface an elongated bilocular synangium attached to a very short stalk ([fig. 120], B). Reference has already been made to the divergent opinions as to the morphological nature of the sporophylls or sporangiophores, but recent investigations distinctly favour the view that a sporophyll is best interpreted as a stalked leaf with two sterile laminae and an almost sessile, or in some cases a more obviously stalked, synangium; the whole sporophyll is characterised by the possession of a ventral and a dorsal lobe[47]. The drawings reproduced in [fig. 120], D and F, illustrate some of the frequent variations described by Thomas in plants which he observed in the New Zealand forests. The sporophyll shown in [fig. 120], D and F, has branched twice and bears three synangia.

Fig. 120. Tmesipteris.

• A. Foliage leaves.
• B. Sporophyll and bilocular synangium.
• C. Diagram of transverse section of stele. px, protoxylem.
• D, F. Abnormal sporophylls. (From drawings made by Prof. Thomas and generously placed at my disposal. A.C.S.)
• E. Portion of C enlarged.

The aerial branches of Tmesipteris possess a central cylinder of separate xylem groups in which the protoxylem occupies an internal position ([fig. 120], C and E, px) enclosing an axial parenchymatous region. The cells of a few layers of the inner cortex immediately outside the endodermis are rendered conspicuous by a dark brown deposit. The cortex as a whole is composed of uniform parenchymatous tissue. In the lower part of the aerial shoots and in the rhizome the xylem forms a solid strand without protoxylem elements and conforms more clearly to that of Psilotum.

In this short account of the anatomy of Tmesipteris no mention is made of the effect produced on the stele by the departure of leaf-traces and of vascular stands to supply branches. Miss Sykes[48] in a recently published paper on the genus has shown that the exit of a leaf-trace does not break the continuity of the xylem of the stele, while the exit of a sporophyll-trace is marked by an obvious gap. Evidence is adduced in support of the conclusion that this difference, which at first sight appears to be one of morphological importance, is in reality merely a question of degree and “is due to the earlier preparation for the formation of ‘sporophyll’ than leaf-traces.” Miss Sykes gives her adherence to the view that the “sporophylls” of Tmesipteris are branches and not leaves, but despite the arguments advanced this interpretation seems to me less probable than that which recognises the sporophyll as a foliar organ. Prof. Lignier[49] has pointed out that if Miss Sykes’s conclusion as to the axial nature of the sporophyll in Tmesipteris is accepted, it diminishes the force of the comparison between the sporophylls of that genus and Sphenophyllum as those of the latter can hardly be regarded as other than foliar organs.

Both members of the Psilotales may, as Boodle has suggested, be regarded as descendants of a common parent in which the aerial stems possessed a fluted or stellate cylinder of mesarch xylem. There can be no doubt as to the significance of the morphological resemblances between the Psilotales and the genera Sphenophyllum and Cheirostrobus, but the position of Tmesipteris and Psilotum in the plant-kingdom may probably be best expressed by adopting the group-name Psilotales rather than by transferring the recent genera to the Sphenophyllales. One of the most striking differences between the Psilotales and the genus Lycopodium is in the form of the sporophylls and sporangia; in Lycopodium a single sporophyll bears a unilocular sporangium, but in the Psilotales the sporophyll may be described as a bilobed structure homologous with a foliage-leaf, bearing a sporangiophore which consists of a short stalk terminating in a bilocular or trilocular synangium; the short stalk receives a special branch from the vascular bundle of the sterile portion of the sporophyll[50].

Fossils described by authors as being closely allied to Psilotum.

A search through palaeobotanical literature reveals the existence of a very small number of specimens which have been identified as representatives of the Psilotales. An inspection of the material or published drawings leads one to the conclusion that practically no information of a satisfactory kind is available in regard to the past history of the two southern genera Psilotum and Tmesipteris, which are regarded by some botanists as relics of an ancient branch[51] of pteridophytes.

PSILOTITES, ETC.

In 1842 Münster[52] instituted the genus Psilotites for a small impression of a slender branched axis from Jurassic rocks near Mannheim in Germany which he named Psilotites filiformis; Schimper[53] spoke of the specimens as too doubtful for determination, an opinion with which every botanist would cordially agree. Goldenberg’s species Psilotites lithanthracis[54] from the Saarbrücken coal-field is founded on impressions of axes: some of these are dichotomously branched and bear small oval projections, which may be rudimentary leaves or possibly leaf-scars. More recently Kidston[55] described specimens of branched axes from the Lanarkshire coal-field bearing a row of lateral thorn-like projections under the title Psilotites unilateralis; but these fragments, as Dr Kidston himself admits, are of no botanical value.

In a paper on fossil Salvinias, Hollick[56] mentions Salvinia reticulata, originally described by Heer and by Ettingshausen and S. Alleni Lesq.[57] a Tertiary species, and calls attention to their very close resemblance in form, nervation, and apex to the leaves of the genus Tmesipteris: he refers both species to that genus. The drawings reproduced by Hollick represent leaves with a midrib and numerous anastomosing lateral veins, whereas in Tmesipteris the lamina of the leaf has a midrib without lateral branches. An enlarged drawing of the outlines of the epidermal cells would correspond closely with the small reticulations in the fossil leaves and it may be that there has been some confusion between veins and cell-outlines. In any case there would seem to be no reason for the use of the recent generic name[58].

Among other fossils assigned to the Psilotales we have Marion’s genus Gomphostrobus from the Permian of France and Germany[59]. Marion placed this plant in the Coniferales on the strength of its resemblance to Walchia and Araucaria, but Potonié[60] is inclined to recognise in the leaves and monospermic sporophylls characters suggestive of Lycopodiaceous affinity.

The latter author in 1891[61], in ignorance of Marion’s proposal to adopt the name Gomphostrobus, instituted a genus Psilotiphyllum for the sporophylls of a species originally described by Geinitz[62] as Sigillariostrobus bifidus, but he subsequently adopted Marion’s designation and with some hesitation included the French and German specimens in the Psilotales. As stated elsewhere[63], Potonié’s arguments in favour of his view hardly carry conviction, and it is probably more in accordance with truth to deal with Gomphostrobus in the chapter devoted to the Coniferales.

Psilophyton.

The generic title Psilophyton, instituted by the late Sir William Dawson[64], has become familiar to geologists as that of a Pre-Carboniferous plant characteristic of Devonian and Silurian rocks in Canada, the United States of America, and Europe. From the botanist’s point of view the name stands for miscellaneous remains of plants of different types and in many cases unworthy of record. The genus was founded on impressions of branched axes from the Devonian strata of New Brunswick resembling the rachis and portions of lateral pinnae of ferns or the forked slender twigs of a Lycopod. The type-species Psilophyton princeps Daws. as represented on somewhat slender evidence in Dawson’s restoration, which accompanies the original description of the genus and has since been copied by several authors, is characterised by the possession of a horizontal rhizome bearing numerous rootlets and giving off dichotomously branched aerial shoots with spinous appendages, compared with rudimentary leaves, and terminating in slender branchlets bearing pendulous oval “spore-cases” from their tips. Some of the branchlets exhibit a fern-like vernation. The plant is spoken of by Dawson as apparently a generalised type[65], resembling in habit and in its rudimentary leaves the recent genus Psilotum and presenting points of contact with ferns. Specimens were found in an imperfectly petrified state showing a central cylinder of scalariform tracheae surrounded by a broad cortical zone of parenchyma and fibrous tissue.

Among other species described by the author of the genus we need only mention Psilophyton robustius, characterised by vegetative shoots and “spore-cases” similar to those of the type-species; but, as Solms-Laubach[66] has pointed out, the petrified sections referred by Dawson to P. robustius are of an entirely different anatomical type from that of P. princeps[67].

British fossils from the Old Red Sandstone from the north of Scotland, Orkney and Caithness, originally figured by Hugh Miller and compared by him with algae but more especially with recent Lycopods, were subsequently placed by Carruthers[68] in the genus Psilophyton as P. Dechianum, the specific designation being chosen on the ground that the Scotch specimens are specifically identical with fossils described by Goeppert[69] as Haliserites Dechianus.

Various opinions have been expressed in regard to the nature of the Devonian species Haliserites Dechianus Goepp. with which Carruthers[70] identified Miller’s Old Red Sandstone plant: reference may be made to a paper by White[71] containing figures of dichotomously branched impressions described as species of Thamnocladus which he includes among the algae.

In describing some Belgian impressions of Devonian age as Lepidodendron gaspianum Daws. Crépin[72] states that Carruthers has come to regard the specimens named by him Psilophyton Dechianum as branches of a Lepidodendron; he also quotes Carruthers as having expressed the opinion that the name Psilophyton had been employed by Dawson for two kinds of fossils, some being twigs of Lepidodendron while others, identified by Dawson as the reproductive branches of species of Psilophyton, represent the spore-cases of ferns comparable with Stur’s genus Rhodea[73]. One of the examples figured by Carruthers[74] as P. Dechianum from Thurso (preserved in the British Museum, no. 52636), measuring 34 cm. in length and 8 mm. broad, bears a close resemblance to a fern rhizome covered with ramental scales such as that of a species of Davallia. Other Belgian specimens described by Gilkinet[75] as Lepidodendron burnotense, like Crépin’s species, are no doubt generically identical with some of the Scotch and Canadian fossils placed in the genus Psilophyton, though Penhallow[76] considers that the species Lycopodites Milleri is more correctly referred to Lycopodites than to Psilophyton.

A more recent paper on the Geology of the Perry basin in South-eastern Maine by Smith and White[77] contains a critical summary of the literature on Psilophyton and drawings of specimens. The latter afford good examples of Pre-Carboniferous plant fragments, such as are often met with in various parts of the world, which conform in habit to the New Brunswick specimens made by Dawson the type of his genus.

An examination of material in the Montreal Museum and of Hugh Miller’s specimens in the Edinburgh collection leads me to share the opinion of Count Solms-Laubach that the name Psilophyton has been applied to plants which should not be included under one generic title. As Kidston[78] pointed out, the Canadian species Psilophyton robustius is not generically distinct from British and Belgian specimens referred to Lepidodendron; it may possibly be identical with the Bohemian plants on which Stur founded his genus Hostinella[79]. The Devonian plants described by Stur have since been examined by Jahn[80] who regards them as vascular plants, and not as algae to which Stur referred them; he mentions two species of Psilophyton but gives no figures.

The “spore-cases” of Dawson may be found to be the microsporangia or perhaps the small seeds of some pteridosperm; the forked axes with a smooth surface and others figured by Miller and by Dawson, with the surface covered with scales suggesting the ramenta of a fern, may be the rachises or rhizomes of filicinean plants. Other specimens may be Lepidodendron twigs, as for example the petrified fragments figured by Dawson as Psilophyton princeps; while the stem identified as P. robustius is most probably that of a Gymnosperm. It is doubtful whether a useful purpose is served by retaining the genus Psilophyton. It was in the first instance instituted on the assumption, which cannot be upheld, that the abundant material in the New Brunswick beds bore a sufficiently close resemblance to the rhizome and aerial branches of Psilotum. Psilophyton has served as a name for miscellaneous plant fragments, many of which are indeterminable. Dr White concludes his account of the genus with the following words[81]:

“The examination of such so-called Psilophyton material as I have seen shows the existence in America of two or more groups, represented by several fairly well-marked species which possess stratigraphical value, and which should be carefully diagnosed and illustrated. It is probable also that additional material throwing light on the structure and relationships of these very remarkable early types of land-plants will be discovered at some locality. The inspection of the material in hand emphasises the need, as was pointed out by Solms-Laubach, for the revision of the material referred by various authors to Psilophyton, together with a thorough re-examination and re-publication of the types.”

Until a thorough re-examination has been made of the Canadian material, with a view to determine whether there exist substantial reasons for the retention of Dawson’s genus, it is undesirable to continue to make use of this name for Pre-Carboniferous fossils which are too incomplete to be assigned with certainty to a definite group of plants. Dr White draws attention to the similarity of some of the Perry basin specimens to Nathorst’s genus Cephalotheca[82] from Devonian rocks of Bear Island in the Arctic regions, a comparison which might be extended to other genera and which serves to illustrate the possibility that many of the specimens labelled Psilophyton may eventually be recognised as examples of well defined generic types belonging to more than one group of plants.

CHAPTER XIV.

LYCOPODIALES.

The recent members of the Lycopodiales are considered apart from the extinct genera in order that our examination of the latter may be facilitated by a knowledge of the salient characteristics of the surviving types of this important section of the Pteridophyta. A general acquaintance with the extinct as well as with the recent genera will enable us to appreciate the contrasts between the living and the fossil forms and to realise the prominent position occupied by this group in the Palaeozoic period, a position in striking contrast to the part played by the diminutive survivors in the vegetation of the present day. In the account of the recent genera special attention is drawn to such features as afford a clue to the interpretation of the fossils, and the point of view adopted, which at times may appear to lead to an excessive attention to details, is necessarily somewhat different from that represented in botanical text-books[83].

A. HOMOSPOREAE.

Lycopodiaceae: genera Phylloglossum, Lycopodium.

B. HETEROSPOREAE.

Selaginellaceae: genus Selaginella.
Isoetaceae: genus Isoetes.

The existing plants included in the Lycopodiales are in nearly all cases perennial herbaceous pteridophytes, exhibiting in their life-histories a well marked alternation of generations. The sporophyte (asexual generation) is characterised by the relatively small size of the leaves except in the genus Isoetes ([fig. 132]) and in the Australian and New Zealand genus Phylloglossum. The stems are usually erect or trailing, pendulous in epiphytic species or small and tuberous in Isoetes and Phylloglossum. The repeated forking of the shoots (monopodial and dichotomous branching) is a prominent feature of the group. The vascular tissue of the stem usually assumes the form of a single axial strand (stele) ([fig. 125]), but the shoots of some species of Selaginella often contain two or more distinct steles ([fig. 131]). The group as a whole is characterised by the centripetal development of the xylem composed almost entirely of scalariform tracheids: secondary xylem and phloem of a peculiar type occur in Isoetes, and the production of secondary xylem elements in a very slight degree has been noticed in one species of Selaginella (S. spinosa)[84]. The roots are constructed on a simple plan, having in most cases only one strand of spiral protoxylem elements (monarch structure). In Lycopodium, in which stem and root anatomy are more nearly of the same type than in the majority of plants, several protoxylem strands may be present. The sporangia are axillary or, more frequently, borne on the upper surface of sporophylls, which are either identical with or more or less distinct from the foliage leaves; in the latter case the sporophylls often occur in the form of a well defined strobilus (cone) at the tips of branches.

The gametophyte (sexual generation) is represented by prothalli which, in the homosporous genera, may live underground as saprophytes, or the upper portion may develop chlorophyll and project above the surface of the ground as an irregularly lobed green structure (e.g. Lycopodium cernuum)[85]. In the heterosporous forms the prothalli are much reduced and do not lead an independent existence outside the spore by the membrane of which they are always more or less enclosed. The sexual organs are represented by antheridia and archegonia; the male cells are provided with two cilia except in Isoetes which has multiciliate antherozoids like those of the ferns.

The existing Lycopods, though widely distributed, never grow in sufficiently dense masses to the exclusion of other plants to form a conspicuous feature in the vegetation of a country. The inconspicuous rôle which they play among the plant-associations of the present era affords a striking contrast to the abundance of the arborescent species in the Palaeozoic forests of the northern hemisphere.

•••••

Lycopodiaceae. Lycopodium, represented by nearly 100 species, forms a constituent of most floras: epiphytic species predominate in tropical regions, while others flourish on the mountains and moorlands of Britain and in other extra-tropical countries. For the most part Lycopodium exhibits a preference for a moist climate and appears to be well adapted to habitats where the amount of sunlight is relatively small and the conditions of life unfavourable for dense vegetation. Mountains and islands constantly recur as situations from which species have been recorded. Some species are essentially swamp-plants, e.g. Lycopodium inundatum, a British species, and L. cruentum from the marshes of Sierra Nevada. A variety of the American species, L. alopecuroides (var. aquaticum) affords an instance of a submerged form, which has been collected from an altitude of 12–14,000 ft. on the Andes and Himalayas. It is noteworthy that a considerable variety of habitats is represented within the limits of the genus and that many species are sufficiently hardy to exist in circumstances which would be intolerable to the majority of flowering plants[86].

The British species frequently spoken of as Club Mosses, include Lycopodium Selago, L. annotinum, L. clavatum, L. alpinum, and L. inundatum.

•••••

Selaginellaceae. The species of Selaginella, over 300 in number, are widely spread in tropical and subtropical forests, growing on the ground with trailing, suberect or erect stems climbing over taller and stouter plants or as pendulous epiphytes on forest trees.

Selaginella lepidophylla, a tropical American type, popularly known as the Resurrection plant, and often erroneously spoken of as the Rose of Jericho[87], possesses the power of rolling up its shoots during periods of drought and furnishes an example of a species adapted to conditions in marked contrast to those which are most favourable to the majority of species.

The only British species is Selaginella spinosa named by Linnaeus Lycopodium selaginoides and occasionally referred to as Selaginella spinulosa A. Br. (not to be confounded with a Javan species S. spinulosa Spring[88]).

•••••

Isoetaceae. Isoetes ([fig. 132]), of which Mr Baker in his Handbook of the Fern-Allies enumerates 49 species, is a type apart, differing in habit as in certain other characters from the other members of the Lycopodiales. Some botanists[89] prefer to include the genus among the Filicales, but the balance of evidence, including resemblances between Isoetes and extinct Lycopodiaceous plants, would seem to favour its retention as an aberrant genus of the group Lycopodiales. Some species are permanently submerged, others occur in situations intermittently covered with water, and a few grow in damp soil. Isoetes lacustris is found in mountain tarns and lakes of Britain and elsewhere in Central and Northern Europe and North America. Isoetes hystrix[90], a land-form occurs in Guernsey, North-East France, Spain and Asia Minor.

Lycopodiaceae.

The monotypic genus Phylloglossum, represented by P. Drummondii of Australia and New Zealand, though interesting from the point of view of its probable claim to be considered the most primitive type of existing Lycopodiaceous plants, need not be dealt with in detail. A complete individual, which does not exceed 4 or 5 cm. in length, consists of a very small tubercle or protocorm bearing a rosette of slender subulate leaves and prolonged distally as a simple naked axis which overtops the foliage leaves and terminates in a compact cluster of small scale-like sporophylls, each subtending a single sporangium[91].

Lycopodium. It would be out of place in a volume devoted mainly to fossil plants to attempt a comprehensive account of the general morphology of recent species, and indeed our knowledge of the anatomical characters of the genus is still somewhat meagre. For purposes of comparison with extinct types, it is essential that some of the more important morphological features of existing species should be briefly considered. The additions made to our knowledge of the gameophyte[92] of European and tropical species during the last two decades have revealed a striking diversity in habit.

In several species, grouped round the widely distributed type Lycopodium Selago Linn., the comparatively short, erect or suberect, shoots form fairly compact tufts; the ordinary foliage-leaves function as sporophylls, and the sporangia are not localised on special portions of shoots. From this type, we pass to others in which the fertile leaves tend to be confined to the tips of branches, but hardly differ in form from the sterile. A further degree of specialisation is exhibited by species with well-defined cones composed of leaves (or bracts), the primary function of which is to bear sporangia and to afford a protective covering to the strobilus[93].

Lycopodium rufescens Hook. An Andian species with stout dichotomously branched erect stems bears on the younger shoots crowded leaves with their thick and broadly triangular laminae pointing upwards, but on the older and thick shoots the laminae are strongly reflexed ([fig. 121], A). The lower part of the specimen represented in [fig. 121], A, shows tangentially elongated scars and persistent leaf-bases or cushions left on the stem after the removal of the free portions of the leathery leaves, a surface-feature which also characterises the Palaeozoic genus Lepidodendron. The reflexed leaves and persistent leaf-cushions are clearly seen in the piece of old stem of Lycopodium dichotomum Jacq., a tropical American species reproduced in [fig. 121], B. Such species as L. erythraeum Spring, and others with stiff lanceolate leaves exhibit a striking resemblance to the more slender shoots of some recent conifers, more especially Araucaria excelsa, A. Balansae, Cryptomeria, Dacrydium and other genera.

Fig. 121. Lycopodium.

1. Lycopodium rufescens.
2. L. dichotomum.
3. L. tetragonum.
4. L. nummularifolium.
5. L. Dalhousianum.
6. L. casuarinoides.
7. L. volubile.

Fig. 122. Lycopodium squarrosum. The branches of the larger shoot terminate in cones. (From a plant in the Cambridge Botanic Garden. Reduced.)

In Lycopodium tetragonum Hook., ([fig. 121], C), a species from the Alpine region of the Andes, the long, pendulous and repeatedly forked branches bear four rows of fleshy ovate leaves and simulate the vegetative characters of certain conifers.

Fig. 123. Lycopodium cernuum.
(From a specimen in the Cambridge Herbarium. ½ nat. size.)

L. squarrosum Forst. ([fig. 122]) a tropical species from India, Polynesia, and other regions, is characterised by its stout stems reaching a diameter of 2·5 cm., bearing long pendulous branches with large terminal cones composed of sporophylls differing but slightly from the foliage leaves. The plant represented in the photograph serves as a good illustration of the practical identity in habit between Palaeozoic and recent genera.

Fig. 124. Lycopodium obscurum.

L. Dalhousianum Spring, from the mountains of the Malay Peninsula and Borneo, has larger leaves of finer texture with a distinct midrib reaching a length of 2–3 cm. ([fig. 121], E). Another type is illustrated by L. nummularifolium Blume, also a Malayan species, in which the leaves are shorter, broadly oblong or suborbicular, and the branches terminate in narrow and often very long strobili (sometimes reaching a length of 30 cm.) with small bracts in striking contrast to the foliage leaves ([fig. 121], D). A similar form of long and slender strobilus occurs in L. Phlegmaria Linn., a common tropical Lycopod: the frequent forking of the strobili noticed in this and other species is a character not unknown among fossil cones (Lepidostrobi).

L. cernuum Linn. ([fig. 123]), another widely spread tropical type, offers an even closer resemblance than L. squarrosum to the fossil Lepidodendra. The stiff erect stem, reaching in some cases a length of several feet, bears numerous repeatedly forked branches, with crowded linear leaves, terminating in short cylindrical cones with broadly ovate sporophylls. A similar habit characterises the North American species L. obscurum Linn. ([fig. 124]) bearing cones several centimetres in length.

L. casuarinoides Spring ([fig. 121], F) an eastern tropical species, is worthy of notice as exhibiting a peculiar form of leaf consisting of a very small lamina, 3 mm. in length, borne on the top of a long decurrent base, which forms a narrow type of leaf-cushion, bearing some resemblance to the long and rib-like cushions of certain species of Sigillaria, and recalling the habit of slender fossil twigs referred to the Coniferae under such names as Widdringtonites, Cyparissidium, Sphenolepidium.

L. volubile Forst. ([fig. 121], G) a New Zealand species, in habit and leaf-form bears a close resemblance to the Jurassic Lycopodites falcatus Lind. and Hutt. ([fig. 137]): it is also a representative of a few species of Lycopodium which agree with the majority of species of Selaginella in having two kinds of sterile leaves, comparatively long falcate leaves forming two lateral rows and smaller appressed leaves on the upper surface of the branches.

These examples suffice to illustrate the general appearance presented by the vegetative shoots of recent species of which the foliage leaves vary considerably—from the small scale-leaves of Lycopodium tetragonum, to the very slender linear subulate leaves of such a species as L. verticillatum Linn. or the long and broader lamina of L. Dalhousianum ([fig. 121], E). It is obvious that fragments of the various types preserved as fossils might well be mistaken either for some of the larger mosses or for twigs of conifers. As Dr Bommer[94] has pointed out in his interesting paper on “Les causes d’erreur dans l’étude des empreintes végétales” some dicotyledonous plants may also simulate the habit of Lycopods: he cites Phyllachne clavigera Hook (Candolleaceae), Tafalla graveolens Wedd (Compositae) and Lavoisiera lycopodioides Gard. (Melastomataceae). Another point illustrated by [fig. 121] is the close agreement in habit and in the form of the leaves and leaf-cushions between the recent plants and the Palaeozoic Lepidodendreae.

In his masterly essay “On the vegetation of the Carboniferous Period, as compared with that of the present day” Sir Joseph Hooker called attention to the variation in the shape and arrangement of the leaves in the same species of Lycopodium. The three woodcuts which he publishes of Lycopodium densum, a New Zealand species, afford striking examples of the diversity in habit and leaf-form and justify his warning “that if the species of Lepidodendron were as prone to vary in the foliage as are those of Lycopodium, our available means for distinguishing them are wholly insufficient[95].”

As we have already noticed, there is a considerable diversity among recent species, both as regards habitat and habit; in the anatomy of the stem also corresponding variations occur within the limits of a well-defined generic type of stele. In species with creeping stems, such as L. clavatum[96], the stele exhibits an arrangement of vascular tissue characteristic of the plagiotropic forms. The xylem consists of more or less horizontal plates of scalariform tracheae, each surrounded by small-celled parenchyma, alternating with bands or groups of somewhat ill-defined phloem. The protoxylem and protophloem elements occupy an external position (exarch), pointing to a centripetal development of the metaxylem. This centripetal or root-like character of the primary xylem is an important feature in recent as in fossil Lycopods. The close agreement between the roots and stems of recent species in the disposition of the vascular elements also denotes a simpler type of anatomy than occurs in the majority of vascular plants in which stem and root have more pronounced structural peculiarities. A pericycle, 2–6 cells in breadth, encloses the xylem and phloem bands and this is succeeded by an endodermis, 2–3 cells broad, with vaguely defined limits. In L. clavatum, as in L. alpinum, another British species, the broad cortex is differentiated into three fairly distinct regions; abutting on the endodermis is a zone several layers broad of thick-walled cells constituting an inner cortex modified for protection and support; the central region consists of larger and thinner-walled cells adapted for water-storage and aeration; beyond this is an outer cortical zone of firmer and thicker elements. The prominent leaf-bases or leaf-cushions ([fig. 125], A, lc) give to the surface of a transverse section a characteristic appearance which presents the closest agreement with that of the younger shoots of Lepidodendron. From the peripheral protoxylem groups small strands of xylem are given off, which follow a steeply ascending course through the cortex to the single-veined leaves. The leaf-traces, in several species at least, are characterised by a mesarch structure ([fig. 125], F, G), the spiral protoxylem elements occupying an approximately central position. The mesophyll of the leaves varies in regard to the extent of differentiation into a palisade and spongy parenchyma; in all cases there is a single vascular bundle occasionally accompanied by a secretory duct.

Fig. 125.

1. Lycopodium dichotomum. Transverse section of stem: lc, leaf-cushion; lt, leaf-trace; R, roots.
2. L. cernuum, portion of cortex of fig. H, enlarged.
3. L. saururus. Cortex: lt, leaf-trace; a, thin-walled tissue; b, thick-walled tissue; lc, lacuna.
4. L. saururus. Stele: x, xylem; p, phloem.
5. Portion of fig. D, enlarged: px, protoxylem; p, phloem.
6. Transverse section of leaf of Lycopodium.
7. Vascular bundle of leaf: px, protoxylem.
8. L. cernuum: b, branch of stele; c–c″, cortex; s, space in cortex; lt, leaf-trace.
9. Stele of fig. H, enlarged (phloem omitted).

In erect stems of Lycopodium, as represented by L. cernuum (figs. [123], [125], H, I), L. Dalhousianum, L. squarrosum ([fig. 122]) and many others, the stele presents a characteristic appearance due to the xylem plates being broken up into detached groups or short uniseriate bands with the interspaces occupied by phloem islands. This type of structure bears a superficial resemblance to that in the single stele of certain species of the fern Lygodium[97], but it is distinguished by the islands of phloem scattered through the stele. In other species the xylem tends to assume the form of a Maltese cross (e.g. L. serratum Thbg.) or it may be disposed as V-shaped and sinuous bands terminating in broad truncate ends composed of protoxylem elements. This form of the xylem and the distribution of the phloem groups are shown in [fig. 125], D, E, drawn from a section of a plant of Lycopodium saururus Lam.[98] collected by Mr A. W. Hill at an altitude of 15,000 feet on the Andes of Peru. The position of the protoxylem is shown [fig. 125], E, px.

While several species possess a cortex of three distinct zones ([fig. 125], H, c, c′, c″), in others the extra-stelar tissue is much more homogeneous, consisting of thin-walled parenchyma or in some cases of thick-walled elements; as a general rule, however, there is a tendency towards a more compact arrangement in the inner and outer portions of the cortex as contrasted with the larger and more loosely connected cells of the middle region. In certain types the middle cortex contains fairly large spaces, as in the swamp-species L. inundatum, which with L. alopecuroides exhibits another feature of some interest first described by Hegelmaier[99]. If a transverse section of the stem of L. inundatum be examined the leaf-traces are seen to be accompanied by a circular canal containing mucilage which extends into the lamina of the leaf. In a specimen of L. cernuum[100] obtained at a height of 2500 ft. by Professor Stanley Gardiner in the Fiji Islands, the leaf-traces ([fig. 125], B lt) were found to be accompanied for part of their course by a well-marked secretory space ([fig. 125], B, s). There is little doubt that the presence of these mucilage canals is directly connected with a certain type of habitat[101] and attention is called to them in view of a resemblance which they offer to a characteristic strand of tissue, known as the parichnos, which is associated with the leaf-traces of Lepidodendreae and Sigillarieae. In the section shown in [fig. 125], H, the xylem of the stele forms more continuous bands than is often the case in L. cernuum which has already been described as having its xylem in small detached groups. The presence of the smaller branch-stele ([fig. 125], H, b) affords an example of monopodial branching. The outer cortex of L. saururus ([fig. 125], C) exhibits a somewhat unusual feature in the distribution of the thicker-walled tissue (b) which encloses a patch of more delicate parenchyma (a) with large lacunae (lc) in the region of the leaf-bases, and presents the appearance of an irregular reticulum. This arrangement of the mechanical tissue in the outer cortex is comparable with that in stems of some species of Sigillaria.

In certain species of Lycopodium the roots[102], which arise endogenously from the axial vascular cylinder, instead of passing through the cortex of the stem by the shortest route, bend downwards and bore their way in a more or less vertical direction before emerging at or near the base of the aerial shoot. The transverse section of L. dichotomum represented in [fig. 125], A, shows several roots (R) in the cortex; they consist of a xylem strand of circular or crescentric form accompanied by phloem and enclosed by several layers of root-cortex. The roots of Lycopodium do not always present so simple a structure as those of L. dichotomum; the xylem may have an irregularly stellate form with as many as ten protoxylem groups.

Reproductive Shoots[103]. In Lycopodium Selago the foliage leaves serve also as sporophylls and, as Professor Bower[104] has pointed out, the branches exhibit to some extent a zonal alternation of sterile and fertile leaves; in other species, in which foliage leaves and sporophylls are practically identical, the sporangia occur sporadically on the ordinary leaves. In species with well-defined terminal cones the lower sporophylls may bear arrested sporangia and thus form transitional stages between sterile and fertile leaves, a feature which occurs also in the male and female flowers of many recent Araucarieae[105]. The sporangia[106] ([fig. 126], D, F) are usually reniform and compressed in a direction parallel to the surface of the cone-scales; they are developed from the upper surface and close to the base of the fertile leaf to which they are attached by a short and thick stalk (e.g. L. inundatum) or by a longer and more slender pedicel (L. Phlegmaria, [fig. 126], E). On maturity the sporangia open as two valves in the plane of compression and the line of dehiscence is determined in some species at least by the occurrence of smaller cells in the wall. In transverse sections of cones in which the sporangia are strongly saddle-shaped, the sporophylls may appear to bear two sporangia. This is well shown in the section of a cone of L. clavatum shown in [fig. 126], F. The sporangia a and b are cut through in an approximately median plane showing the irregular outline of the sterile pad (p) of tissue in the sporogenous cavity. Those at c and d have been traversed at a lower level and the two lobes of the saddle-shaped sporangia are cut below the attachment to the sporophyll. The distal laminae of the sporophylls, cut at different levels, are seen at the periphery of the cone.

Fig. 126.

1. Lycopodium cernuum, longitudinal section of strobilus; a, band of lignified cells.
2. L. cernuum. Cell from sporangium wall.
3. L. cernuum. Sporophyll and sporangium; lt, vascular bundle.
4. L. clavatum. Part of radial longitudinal section of strobilus; p, sterile tissue.
5. L. Phlegmaria. Sporophyll and stalked sporangium.
6. L. clavatum. Transverse section of strobilus; p, sterile pad.

In longitudinal radial section of some cones the sporangia appear to occupy an axillary position, but in others (e.g. L. clavatum) they are attached to the horizontal portion of the sporophyll almost midway between the axis of the cone and the upturned distal end of the sporophyll ([fig. 126], D). The wall of a sporangium frequently consists of 2–3 cell-layers and in some cases (e.g. L. dichotomum), it may reach a thickness of seven layers, resembling in this respect the more bulky sporangia of a certain type of Lepidodendroid cone. The sporogenous tissue is separated from the stalk of the sporangium by a mass of parenchymatous tissue which may project as a prominent pad ([fig. 126], D, F, p) into the interior of the sporogenous cavity. This basal tissue (the subarchesporial pad of Bower[107]) has been observed in L. clavatum to send up irregular processes of sterile cells among the developing spores, suggesting a comparison with the trabeculae which form a characteristic feature of the sporangia of Isoetes and with similar sterile strands noticed by Bower[108] in Lepidostrobus (cone of Lepidodendron).

Each sporophyll is supplied by a single vascular bundle which according to published statements never sends a branch to the sporangium base. The fertile tips of the foliage shoots of L. cernuum ([figs. 126], A–C) afford good examples of specialised cones. The surface of the cone is covered by the broadly triangular laminae of sporophylls ([fig. 126], C) which in their fimbriate margins resemble the Palaeozoic cone-scales described by Dr Kidston[109] as Lepidostrobus fimbriatus. The distal portions of the sporophylls are prolonged downwards ([fig. 126], A) to afford protection to the lower sporangia, their efficiency being increased by the lignified and thicker walls (A, a) of the cells in the lower portion of the laminar expansion. The cells of the sporangial wall are provided with strengthening bands which in surface-view ([fig. 126], B) present the appearance of prominent pegs. Since the appearance of Miss Sykes’s paper on the sporangium-bearing organs of the Lycopodiaceae, Dr Lang[110] has published a more complete account of the structure of the strobilus of Lycopodium cernuum in which he records certain features of special interest. The importance of these morphological characters is increased by their agreement, as shown by Lang, with those of the Palaeozoic cone Spencerites[111]. The sporophylls of a cone (12 mm. long by 3 mm. in diameter) of Lycopodium cernuum show an abrupt transition from the foliage leaves, but like these they occur in alternate whorls of five. A large sporangium is attached to the upper face of each sporophyll close to the base of the obliquely vertical distal lamina ([fig. 127]); each sporophyll, which is supplied with a single vascular bundle, has a large mucilage-cavity (m) in its lower region. “The mucilaginous change” in the sub-sporangial portion of a sporophyll “extends to the surface involving the epidermis, so that this portion of the sporophyll-base may be described as consisting of a mass of mucilage bounded below by a structureless membrane[112].” Dehiscence of the sporangia occurs at the middle of the distal face ([fig. 127], x). As seen in the radial section ([fig. 127], ma) the outer margin of the base of the sporophyll bears a short outgrowth. The leaf-bases of each whorl hang down between the sporangia of the alternating whorl below, and the base of each sporophyll is coherent with the margins of the two sporophylls of the next lower whorl between which it lies, the sporangia being thus closely packed and lying in a pocket “open only on the outer surface of the cone.” Fig. 128 represents a transverse section through a cone in the plane AA of [fig. 127]; this traverses the sporangia and their subtending bracts (b) of one whorl and the dependent bases of the sporophylls of the next higher whorl in the region of the mucilage-sacs (m), which are bounded at the periphery by the outer tissue of the sporophylls (a). A transverse section in the plane BB of [fig. 127] is shown in [fig. 129]: the pedicels and a part of each vascular strand are seen at b radiating from the axis of the cone; one sporophyll (sp, a) is cut through in the region of the pad of tracheal tissue that characterises the short sporangial stalks. The upper portions of the sporangia of the next lower whorl, which project upwards against the mucilaginous bases of the sporophylls above (cf. [fig. 127], BB) are shown at c and external to them, at a, the section has cut through the outer persistent portions of these sporophyll bases.

Fig. 127. Radial longitudinal section of the cone of Lycopodium cernuum. (After Lang.)

Fig. 128. Transverse section of the cone of Lycopodium cernuum, in its plane AA of [fig. 127]. (After Lang.)

Fig. 129. Transverse section of the cone of Lycopodium cernuum in the plane BB of [fig. 127]. (After Lang.)

As Lang points out, this highly complex structure is an expression of the complete protection afforded to the sporangia of a plant met with in exposed situations in the tropics; it is also of importance from a morphological standpoint as exhibiting an agreement with the extinct type of Lycopod cone represented by Spencerites.

Selaginellaceae.

Selaginella differs from Lycopodium in the production of two kinds of spores, megaspores and microspores, and, in the great majority of species, in the dimorphic character of the foliage leaves, which are usually arranged in four rows, the laminae of the upper rows being very much smaller than those of the lower ([fig. 130], 1–3). The smaller leaves are shown more clearly in [fig. 130], 1a. It is obvious from an examination of a Selaginella shoot, such as is shown in [fig. 130], that in fossil specimens it would often be almost impossible to recognise the existence of two kinds of leaves. Some species, e.g. Selaginella spinosa[113], the sole British representative of the genus, are homophyllous and agree in this respect with most species of Lycopodium. Another feature characteristic of Selaginella, as contrasted with Lycopodium, is the presence of a ligule in both foliage leaves and sporophylls. This is a colourless thin lamina attached by a comparatively stout foot to the base of a pit on the upper surface and close to the lower edge of the leaf ([fig. 130], 4, l; [fig. 131], E, F, l).

Fig. 130. Selaginella grandis. (1–3, nat. size.)

In an erect species, such as S. grandis Moore[114] ([fig. 130] and [fig. 131], G) from Borneo, the main shoots, which may attain a height of 2–3 feet, bear small and inconspicuous leaves of one kind, but the lateral and repeatedly forked shoots are heterophyllous. The passage from the homophyllous to the heterophyllous arrangement is shown in the transition from the erect to the dorsiventral habit of the lateral shoots ([fig. 130], 2). The monopodially or dichotomously branched shoots produce long naked axes at the forks; these grow downwards to the ground where they develop numerous dichotomously forked branches. For certain reasons these naked aerial axes were named rhizophores and have always been styled shoots, the term root being restricted to repeatedly forked branches which the rhizophores produce in the soil. It has, however, been shown by Professor Harvey-Gibson[115] that there is no sufficient reason for drawing any morphological distinction between rhizophores and roots, the term root being applicable to both.

Our knowledge of the anatomy of Selaginella, thanks chiefly to the researches of Harvey-Gibson[116], is much more complete than in the case of Lycopodium. The stems, which may be either trailing or erect, are usually dorsiventral, and it is noteworthy that different shoots of the same plant or even the same axis in different regions may exhibit considerable variation in the structure and arrangement of the vascular tissue. In the well-known species, Selaginella Martensii, the stem, which is partly trailing, partly ascending, possesses a single ribbon-shaped stele composed of scalariform tracheids with two marginal protoxylems formed by the fusion of the leaf-traces of the dorsal and ventral leaves respectively. As in Lycopodium the metaxylem tracheae are as a rule scalariform, but reticulate xylem elements are by no means unknown. The tracheal band, surrounded by parenchymatous elements, is enclosed by phloem with external protophloem elements. The characteristic features of the stele are shown in the diagrammatic drawing of a section of another species—S. Willdenowii—represented in [fig. 131], A.

Fig. 131.

1. Selaginella Willdenowii. Transverse section of stem: a, outer cortex; p, phloem; t, trabeculae.
2. S. spinosa, stem: px, protoxylem.
3. S. laevigata var. Lyallii, section of stele: t, ridge of xylem cylinder; e, endodermis.
4. S. rupestris, seedlings with cotyledons (c) protruding beyond the sporophylls (b).
5. Transverse section of Selaginella leaf-base: l, ligule; lt, leaf-trace.
6. Portion of G. enlarged.
7. S. grandis. Longitudinal section of strobilus: bb, sporophyll-trace; l, ligule.

A pericycle composed of one or two layers of chlorophyll-containing cells encircles the whole stele which is suspended in a lacuna by trabeculae ([fig. 131], A, B, t) connecting the pericycle with the inner edge of the broad cortex. The trabeculae consist in part of endodermal cells characterised by cuticular bands. The cortex is usually differentiated into three fairly distinct regions. Mechanical tissue of thick-walled fibres constitutes the outer region (a); the middle cortex consists of thinner-walled parenchyma, the elements of which become smaller and rather more compactly arranged in the inner zone. The middle cortex is frequently characterised by the presence of spaces and by the hyphal or trabecular structure of the tissue, a feature which, as Bower[117] pointed out, is common to many recent and fossil members of the Lycopodiales. In some cases, e.g. S. erythropus, from tropical America, the cortex of the creeping stem consists entirely of thick-walled cells. Selaginella grandis ([fig. 130]) has “a short decumbent stem rooted at close intervals[118],” from which thick erect aerial shoots rise to a height of one foot or more. In the apical region these erect axes give off repeatedly forked foliage shoots on which the spiral phyllotaxis of the homophyllous axis is gradually replaced by four rows of two kinds of leaves ([fig. 130], 2). The anatomy of this species agrees with that of S. Martensii. The trailing or semi-erect and homophyllous shoots of Selaginella spinosa[119] present a distinct type of vascular anatomy. The upper part of the ascending stem has an axial strand of xylem with seven peripheral groups of spiral protoxylem tracheae ([fig. 131], B); in the trailing portion of the shoot the protoxylem elements occur as one central group in the solid rod of metaxylem through which the leaf-traces pass on their way to the axial protoxylem. This type is important as affording an exception, in the endarch structure of the xylem, to the usual exarch plan of the stelar tissues. This species is the only one in which any indication of the production of secondary xylem elements has so far been recorded. Bruchmann[120] has shown that, in the small tuberous swelling which occurs at the base of the young shoot (hypocotyl), a meristematic zone is formed round the axial vascular strand and by its activity a few secondary tracheids are added to the primary xylem. With this exception Selaginella appears to have lost the power of secondary thickening, the possession of which constitutes so striking a feature of the Palaeozoic Lycopods. Another type is represented by S. inaequalifolia, an Indian species, the shoots of which may have either a single stele or as many as five, each in its separate lacuna. The homophyllous S. laevigata var. Lyallii Spr., a Madagascan species, affords a further illustration of the variation in plan of the vascular tissues within the genus. There is a considerable difference in structure between the erect and creeping shoots; in the former there may be as many as 12–13 steles, which gradually coalesce before the vertical axis joins the creeping rhizome to form one central and four peripheral steles. In the rhizome there is usually a distinct axial stele without protoxylem, surrounded by an ill-defined lacuna and enclosed by a cylindrical stele (solenostele)[121] usually two tracheae in width with four protoxylem strands on its outer edge. The continuity of the tubular stele is broken and, in transverse section, it assumes the form of a horse-shoe close to the base of an erect shoot to which a crescentic vascular strand is given off. Harvey-Gibson[122] has figured a section of the rhizome of this type in which the axial vascular strand is represented by a slight ridge of tracheae ([fig. 131], C, t) projecting towards the centre of the axis of the tubular stele. The cylindrical stele consists of xylem with external and internal phloem (p): cuticularised endodermal cells occur at e and e.

Reference has already been made to the descending naked branches given off from the points of ramification of the foliage shoots of Selaginella. It has been shown by Harvey-Gibson[123] that these branches, originally designated rhizophores by Nägeli and Leitgeb, as well as the dichotomously branched roots which they produce below the level of the ground, possess a single vascular strand of monarch type. It is interesting to find that in some species the aerial portion of the rhizophore has a xylem strand with a central protoxylem, an instance of endarch structure like that in certain portions of the shoot-system of S. spinosa. The root-anatomy of Selaginella and the dichotomous habit of branching afford points of agreement with the subterranean organs of Lepidodendron and Sigillaria.

Leaves. The leaves of Selaginella[124] usually consist of a reticulum of loosely arranged cells, but in some cases part of the mesophyll assumes the palisade form. The single vascular bundle consists of a few small annular or spiral tracheae and at the apex of the lamina the protoxylem elements are accompanied by several short reticulated pitted elements. Both foliage leaves and sporophylls are characterised by the possession of a ligule, a structure which may present the appearance of a somewhat rectangular plate ([fig. 130], 4, l, and [fig. 131], E–G, l) or assume a fan-shaped form with a lobed or papillate margin. The base, composed of large cells, is sunk in the tissue of the leaf close to its insertion on the stem ([fig. 131], E, l) and enclosed by a well-marked parenchymatous sheath. The sheath is separated from the vascular bundle of the leaf by one or more layers of cells, and in some species these become transformed into short tracheids. The ligule is regarded by Harvey-Gibson[125] as a specialised ramentum which serves the temporary function of keeping moist the growing-point and young leaves.

Cones. The terminal portions of the branches of Selaginella usually bear smaller leaves of uniform size which function as sporophylls, but in this genus the fertile shoots do not generally form such distinct cones as in many species of Lycopodium. In S. grandis (figs. [130], 3; [131], G) the long and narrow strobili consist of a slender axis bearing imbricate sporophylls in four rows: each sporophyll subtends a sporangium situated between the ligule and the axis of the shoot. The sporangium may be developed from the axis of the cone or, as in Lycopodium, from the cells of the sporophyll[126]. In some species the lower sporophylls bear only megasporangia, each normally containing four megaspores, the microsporangia being confined to the upper part of the cone. This distribution of the two kinds of sporangia is, however, by no means constant[127]: in some cases, e.g. S. rupestris, cones may bear megasporangia only, and in the cone of S. grandis, of which a small piece is represented in [fig. 131], G, all the sporangia were found to contain microspores.

The occurrence of two kinds of spores in Selaginella constitutes a feature of special importance from the point of view of the relationship between the Phanerogams, in which heterospory is a constant character, and the heterosporous Pteridophytes. One of the most striking distinctions between the Phanerogams and the rest of the vegetable kingdom lies in the production of seeds. Recent work has, however, shown that seed-production can no longer be regarded as a distinguishing feature of the Gymnosperms and Angiosperms. Palaeozoic plants which combined filicinean and cycadean features resembled the existing Phanerogams in the possession of highly specialised seeds. This discovery adds point to the comparison of the true seed with structures concerned with reproduction in seedless plants, which in the course of evolution gave rise to the more efficient arrangement for the nursing, protection, and ultimate dispersal of the embryo. In the megaspore of Selaginella we have, as Hofmeister was the first to recognise in 1851, a structure homologous with the embryo-sac of the Phanerogam. The embryo-sac consists of a large cell produced in a mass of parenchymatous tissue known as the nucellus which is almost completely enclosed by one or more integuments. Fertilisation of the egg-cell within the embryo-sac takes place as a rule while the female reproductive organ is still attached to the parent-plant and separation does not occur until the ovule has become the seed.

In a few cases, notably in certain plants characteristic of Mangrove swamps, continuity between the seed and its parent is retained until after germination. The megasporangium of Selaginella dehisces[128] along a line marked out by the occurrence of smaller cells over the crest of the wall. It has been customary to describe the megaspores as being fertilised after ejection from the sporangia. This earlier separation from the parent and the absence of any protective covering external to the spore-wall constitute two distinguishing features between seeds and megaspores. In Selaginella apus, a Californian species, Miss Lyon has shown that fertilisation of the egg-cell usually takes place while the megaspore is still in the strobilus. On examining withered decayed strobili of this species which had been partially covered with the soil for some months after fertilisation of the megaspores, several young plants were found with cotyledons and roots projecting through the crevices of the megasporangia[129]. From this, adds Miss Lyon, “it seems safe to assume that an embryo may have two periods of growth separated by one of quiescence quite comparable to those of seed plants with marked xerophilous features.”

In another Western American species S. rupestris described by the same writer the cotyledons of young plants were found protruding from the imbricate sporophylls of a withered cone ([fig. 131], D). This species is interesting also from the occasional occurrence of one instead of four megasporangia in a sporangium; a condition which affords another connecting link between the heterosporous Pteridophytes, on the one hand, and the seed-bearing Phanerogams in which the occurrence of a single embryo-sac (megaspore) in each ovule is the rule. The cones of Selaginella rupestris retain connexion with the plant through the winter and fertilisation occurs in the following spring. After the embryo has been formed the megasporangium “becomes sunken in a shallow pit formed by the cushion-like outgrowth of the sporophyll around the pedicel.” It is suggested that this outgrowth may be comparable with the integument which grows up from the sporophyll in the fossil genus Lepidocarpon[130] and almost completely encloses the sporangium. In the drawings given by Miss Lyon no features are recognisable which afford a parallel to the integument of Lepidocarpon. I have, however, endeavoured to show, by a brief reference to this author’s interesting account of the two Californian species, that the physiological and morphological resemblances between the megasporangia of Selaginella and the integumented ovules of the seed-bearing plants are sufficiently close to enable us to recognise possible lines of advance towards the development of the true seed.

Professor Campbell[131] records an additional example of a Selaginella—probably S. Bigelovii—from the dry region of Southern California in which the spores become completely dried up after the embryo has attained some size, remaining in that state until the more favourable conditions succeeding the dry season induce renewed activity.

Isoetaceae.

The genus Isoetes is peculiar among Pteridophytes both in habit and in anatomical features. In its short and relatively thick tuberous stem, terminating in a crowded rosette of subulate leaves like those of Juncus and bearing numerous adventitious roots, Isoetes presents an appearance similar to that of many monocotyledonous plants. The habit of the genus is well represented by such species as Isoetes lacustris and I. echinospora[132] ([fig. 132]) both of which grow in freshwater lakes in Britain and in other north European countries. The latter species bears leaves reaching a length of 18 cm. The resemblance in habit between this isolated member of the Pteridophytes and certain Flowering plants, although in itself of no morphological significance, is consistent with the view expressed by Campbell that Isoetes may be directly related to the Monocotyledons[133].

Fig. 132. Isoetes echinospora (After Motelay and Vendryès).

1. Stem of I. lacustris.
2. Base of sporophyll: l, ligule; spg, sporangium partially covered by velum.

There is as a rule little or no difference between the foliage leaves and sporophylls; in I. lacustris the latter are rather larger and in the terrestrial species I. hystrix[134] the sterile leaves are represented by the expanded basal portions only, which persist like the leaf-bases of Lepidodendron as dark brown scales to form a protective investment to the older part of the stem. The innermost leaves are usually sterile; next to these are sporophylls bearing megasporangia, and on the outside are the older sporophylls with microsporangia. The long and slender portion of the leaf becomes suddenly expanded close to its attachment to the stem into a broad base of crescentic section which bears a fairly conspicuous ligule (figs. [132], B, l, [133], E, l) inserted by a foot or glossopodium in a pit near the upper part of the concave inner face. The ligule is usually larger than that of Selaginella, though of the same type. The free awl-like lamina contains four large canals bridged across at intervals by transverse diaphragms, and in the axial region a single vascular bundle of collateral structure. Other vascular elements, in the form of numerous short tracheids occur below the base of the transversely elongated ligule.

Stomata are found on the leaves of I. hystrix, I. Boryana[135], and in other species which are not permanently submerged. Both microsporangia and megasporangia are characterised by their large size and by the presence of trabeculae or strands of sterile tissue ([fig. 133], E, H, t) completely bridging across the sporangial cavity or extending as irregular ingrowths among the spore-producing tissue. Similar sterile bands, though less abundant and smaller, are occasionally met with in the still larger sporangia of Lepidostrobus; these may be regarded as a further development of the prominent pad of cells which projects into the sporangial cavity in recent species of Lycopodium ([fig. 126], D, p). The sporangia are attached by a very short stalk to the base of a large depression in the leaf-base below the ligule, from the pit of which they are separated by a ridge of tissue known as the saddle, and from this ridge a veil of tissue (the velum) extends as a roof over the sporangial chamber ([fig. 133], E, v). In most species there is a large gap between the lower edge of the velum and that of the sporangial pit, but in I. hystrix this protective membrane is separated from the base of the leaf by a narrow opening, the resemblance of which to the micropyle of an ovule suggested to one of the older botanists the employment of the same term[136]. Mr T. G. Hill[137] has called attention to the presence of mucilage canals in the base of the sporophylls of I. hystrix, which he compares with the strands of tissue known as the parichnos accompanying the leaf-traces of Lepidodendron and Sigillaria in the outer cortex of the stem. The transverse section shown in [fig. 133], H and I, shows two of these mucilage canals in an early stage of development; a strand of parenchymatous elements distinguished by their partially disorganised condition and more deeply stained membranes ([fig. 133], I) runs through the spandrels of the sporophyll tissue close to the upper surface. There is a close resemblance between the structure of these partially formed mucilage-canals and the tissue which has been called the secretory zone in Lepidodendron stems. Fig. 133, H, also shows a large microsporangium with prominent trabeculae (t) lying below the velum. A longitudinal section ([fig. 133], E) through a sporophyll-base presents an appearance comparable with that of an Araucarian cone-scale with its integumented ovule and micropyle. The megaspores are characterised by ridges, spines, and other surface-ornamentation[138]. Though usually unbranched, the perennial stem of Isoetes ([fig. 132]) has in rare cases been found to exhibit dichotomous branching, a feature, as Solms-Laubach[139] points out, consistent with a Lycopodiaceous affinity. The apex is situated at the base of a funnel-shaped depression. The stem is always grooved; in some species two and in others three deep furrows extend from the base up the sides of the short and thick axis towards the leaves: from the sides of these furrows numerous slender roots are given off in acropetal succession. A stele of peculiar structure occupies the centre of the stem; cylindrical in the upper part ([fig. 133], A), it assumes a narrow elliptical or, in species in which there are three furrows, a triangular form in the lower portion of the tuberous stem.

ISOETES

The stem of I. lacustris represented in [fig. 132], A, from which the laminae of the leaves have been removed from the summit affords an example of a species with two furrows. The drawing shows the widely gaping sides of the broad furrow with circular root-scars and a few simple and dichotomously branched roots. A short thick column of parenchymatous tissue projects from a slightly eccentric position on the base of the stem.

Fig. 133. Isoetes lacustris.

• A. Transverse section of stem: cr, cortex; x, x² xylem; c, cambium; a, thin-walled tissue; lt, leaf-traces; b, dead tissue.
• B, C, D. Portions of A enlarged.
• E. Longitudinal radial section of sporophyll-base: v, velum; l, ligule; bb; vascular bundle; m, megaspores; t, sterile tissue.
• F. Longitudinal section through the base of a root.
• G. Transverse section of root.
• H. Transverse section of sporophyll, showing sporangium with trabeculae, t; leaf-trace, (lt), and two groups of secretory cells.
• I. A group of secretory cells enlarged.

The primary vascular cylinder[140] consists of numerous spiral, annular or reticulate tracheids ([fig. 133], A, x) which are either isodiametric or longer in a horizontal than in a vertical direction, associated with parenchyma. Lower in the stem crushed and disorganised xylem elements are scattered through a still living trabecular network of parenchymatous tissue. From the axial cylinder numerous leaf-traces ([fig. 133], A, lt) radiate outwards, at first in a horizontal direction and then gradually ascending towards the leaves. The vascular cylinder is of the type known as cauline; that is, some of the xylem is distinct in origin from that which consists solely of the lower ends of leaf-traces. As in Lycopodium the development of the metaxylem is centripetal.

Von Mohl[141], and a few years later Hofmeister[142], were the first botanists to give a satisfactory account of the anatomy of Isoetes but it is only recently[143] that fresh light has been thrown upon the structural features of the genus the interest of which is enhanced by the many points of resemblance between the recent type and the Palaeozoic Lepidodendreae. A striking anatomical feature is the power of the stem to produce secondary vascular and non-vascular tissue; the genus is also characterised by the early appearance of secondary meristematic activity which renders it practically impossible to draw any distinct line between primary and secondary growth. A cylinder of thin-walled tissue ([fig. 133], A, a) surrounds the primary central cylinder and in this a cambial zone, c, is recognised even close to the stem-apex; this zone of dividing cells is separated from the xylem by a few layers of rectangular cells to which the term prismatic zone has been applied. The early appearance of the cambial activity on the edge of the vascular cylinder is shown in [fig. 133], C, which represents part of a transverse section of a young stem. A leaf-trace, lt, is in connexion with the primary xylem, x′, which consists of short tracheids, often represented only by their spiral or reticulately thickened bands of lignified wall, and scattered parenchyma. Some of the radially elongated cells on the sides of the leaf-trace are seen to be in continuity on the outer edge of the stele, at st, with flattened elements, some of which are sieve-tubes. The position of a second leaf-trace is shown at lt′. External to the sieve-tubes the tissue consists of radially arranged series of rectangular cells, some of which have already assumed the function of a cambium (c). The tissue produced by the cambium on its inner edge consists of a varying amount of secondary xylem composed of very short spiral tracheids; a few of these may be lignified ([fig. 133], A, x²) while others remain thin.

Phloem elements, recognisable by the presence of a thickened reticulum enclosing small sieve-areas (fig 133, B, s) are fairly abundant, and for the rest this intracambial region is composed of thin-walled parenchyma. In longitudinal section these tissues present an appearance almost identical with that observed in a transverse section. Fig. 133, B represents a longitudinal section, through the intracambial zone and the edge of the stele, of a younger stem than that shown in [fig. 133], A. Most of the radially disposed cells internal to the meristematic region are parenchymatous without any distinctive features; a few scattered sieve-tubes (s) are recognised by their elliptical sieve-areas and an occasional tracheid can be detected. The cambium cuts off externally a succession of segments which constitute additional cortical tissue ([fig. 133], A, cr) of homogeneous structure, composed of parenchymatous cells containing starch and rich in intercellular spaces. As the stem grows in thickness the secondary cortex reaches a considerable breadth and the superficial layers are from time to time exfoliated as strips of dead and crushed tissue ([fig. 133], A, b). The diagrammatic sketch reproduced in [fig. 133], A, serves to illustrate the arrangement and relative size of the tissue-regions in an Isoetes stem. In the centre occur numerous spirally or reticulate tracheae scattered in parenchymatous tissue which has been considerably stretched and torn in the peripheral region of the stele; the radiating lines mark the position of the leaf-traces (lt) in the more horizontal part of their course. The zone between the cambium (c) and the edge of the central cylinder consists of radially disposed secondary tissue of short, and for the most part unlignified, elements including sieve-tubes and parenchyma; the secondary xylem elements consist largely of thin-walled rectangular cells with delicate spiral bands, but discontinuous rows of lignified tracheae (x²) occur in certain regions of the intracambial zone. The rest of the stem consists of secondary cortex (cr) with patches of dead tissue (b) still adhering to the irregularly furrowed surface. The structure of the cambium and its products is shown in the detailed drawing reproduced in [fig. 133], D. Many of the elements cut off on the inner side of the cambium exhibit the characters of tracheids: most of these are unlignified, but others have thicker and lignified walls (tr).

I. hystrix appears to be exceptional in retaining its leaf-bases, which form a complete protective investment and prevent the exfoliation of dead cortex. Each leaf-trace consists of a few spiral tracheids accompanied by narrow phloem elements directly continuous with the secondary phloem of the intracambial zone. Dr Scott and Mr Hill have pointed out that a normal cambium is occasionally present in the stem of I. hystrix during the early stages of growth; this gives rise to xylem internally. The few phloem elements observed external to the cambium may be regarded as primary phloem, a tissue not usually represented in an Isoetes stem[144]. The occasional occurrence of this normal cambium, may, as Scott and Hill suggest, be a survival from a former condition in which the secondary thickening followed a less peculiar course. The lower leaf-traces become more or less obliterated as the result of the constant increase in thickness of the broad zone of secondary tissues through which they pass.

The adventitious roots are developed acropetally and arranged in parallel series on each side of the median line of the two or three furrows. The three arms of the triangular stele of I. hystrix and the two narrow ends of the long axis of the stele of I. lacustris, which in transverse section has the form of a flattened ellipse, are built up of successive root-bases. A root of Isoetes ([fig. 133], G) possesses one vascular bundle, x, with a single strand of protoxylem, px, thus agreeing in its monarch structure with the root-bundle in Selaginella and many species of Lycopodium. The cortical region of the root consists of a few layers of outer cortex succeeded by a large space, formed by the breaking down of the inner cortical tissue, into which the vascular bundle projects ([fig. 133], F). The peculiarity of the roots in having a hollow cortex and an eccentric vascular bundle was noticed by Von Mohl[145]. In the monarch bundles, as in the fistular cortex and dichotomous branching, the roots of Isoetes present a striking resemblance to the slender rootlets of the Palaeozoic Stigmaria (see [page 246]). The longitudinal section through the base of a root of Isoetes lacustris shown in [fig. 133], F, affords a further illustration of certain features common to the fossil and recent types.

FOSSIL LYCOPODIALES.

Isoetaceae

The geological history of this division of the Pteridophyta is exceedingly meagre, a fact all the more regrettable as it is by no means improbable that in the surviving genus Isoetes we have an isolated type possibly of considerable antiquity and closely akin to such extinct genera as Pleuromeia and Sigillaria. If Saporta’s Lower Cretaceous species Isoetes Choffati[146], or more appropriately Isoetites Choffati, is correctly determined, it is the oldest fossil member of the family and indeed the most satisfactory among the more than doubtful species described as extinct forms of Isoetes.

Isoetites.

The generic name Isoetites was first used by Münster[147] in the description of a specimen, from the Jurassic lithographic slates of Solenhofen in Bavaria, which he named Isoetites crociformis. The specific name was chosen to express a resemblance of the tuberous appearance of the lower part of the imperfectly preserved and indeterminable fossil to a Crocus corm.

Impressions of Isoetes-like leaves from the Inferior Oolite of Yorkshire figured by Phillips[148] and afterwards by Lindley[149] as Solenites Murrayana were compared by the latter author with Isoetes and Pilularia, but these leaves are now generally assigned to Heer’s gymnospermous genus Czekanowskia. An examination of the structure of the epidermal cells of these Jurassic impressions convinced me that they resemble recent coniferous needles more closely than the leaves of any Pteridophyte. The genus Czekanowskia[150] is recognised by several authors as a probable member of the Ginkgoales.

Isoetites Choffati. Saporta.

The late Marquis of Saporta founded this species on two sets of impressions from the Urgonian (Lower Cretaceous) of Portugal which, though not found in actual organic connexion, may possibly be portions of the same plant. Small relatively broad tuberous bodies reaching a breadth of 1 cm. are compared with the short and broad stem of Isoetes, which they resemble in bearing numerous appendages radiating from the surface like the roots of the recent species; on the exposed face of the stem occur scattered circular scars representing the position of roots which were detached before fossilisation. Other impressions are identified as the basal portions of sporophylls bearing sporangia: these suggest the expanded base of the fertile leaves of Isoetes with vertically elongated sporangia, some of which have a smooth surface while in others traces of internal structure are exposed; the interior consists of an irregular network with depressions containing carbonised remains of spores.

While recognising a general resemblance to the sporophylls of Isoetes, certain differences are obvious: there is no ligule in the fossil leaves nor are there any distinct traces of vascular strands such as occur in the leaves of recent species. The form of the sporangium, more elongated than in the majority of recent forms, is compared by Saporta with that in a south European species Isoetes setacea Spr.

Such evidence as we have lends support to the inclusion of these Portuguese fossils in the genus Isoetites, but apart from the fact that we have no proof of any connexion between the stems and supposed sporophylls, the resemblance of the latter to those of Isoetes is, perhaps, hardly sufficient to satisfy all reasonable scepticism.

The generic name Isoetopsis was used by Saporta as more appropriate than Isoetes for some Eocene fossils from Aix-en-Provence which are too doubtful to rank as trustworthy evidence of the existence of the recent genus. The species, Isoetopsis subaphylla[151] is founded on impressions of small scales, 4 mm. long, bearing circular bodies which are compared with sporangia or spores.

Other records of fossils referred to Isoetes need not be described as they have no claim to be regarded as contributions towards the past history of the genus. Heer’s Miocene species Isoetites Scheuzeri and I. Braunii Unger[152] from Switzerland are based on unsatisfactory material and are of no importance.

Pleuromeia.

The generic name Pleuromeia, was suggested by Corda[153] for a fossil from the Bunter Sandstone, the original description of which was based by Münster[154] on a specimen discovered in a split stone from the tower of Magdeburg Cathedral.

The majority of the specimens have been obtained from the neighbourhood of Bernburg, but a few examples are recorded from Commern and other German localities: all are now included under the name Pleuromeia Sternbergi. Germar, who published one of the earlier accounts of the species, states that Corda dissented from Münster’s choice of the name Sigillaria and proposed the new generic title Pleuromeia. One of the best descriptions of the genus we owe to Solms-Laubach[155] whose paper contains references to earlier writers. Illustrations have been published by Münster, Germar[156], Bischof[157], Solms-Laubach and Potonié[158].

Pleuromeia Sternbergi. (Münster.)
[Fig. 134].

• 1842. Sigillaria Sternbergii, Münster.
• 1854. Sagenaria Bischofii, Goeppert[159].
• 1885. Sigillaria oculina, Blanckenhorn.
• 1904. Pleuromeia oculina, Potonié.

Pleuromeia Sternbergi is represented by casts of vegetative and fertile axes, but the preservation of the latter is not sufficiently good to enable us to draw any very definite conclusions as to the nature of the reproductive organs. Casts of the stems reach a length of about 1 metre and a diameter of 5–6 cm., or in some cases 10 cm.; all of them are in a more or less decorticated state, the degree of decortication being responsible for differences in the external features which led Spieker[160] to adopt more than one specific name.

[Fig. 134], A, represents a sketch, made some years ago, of a specimen in the Breslau Museum which contains several examples of this species, among others those described by Germar in 1852. The cylindrical cast (38 cm. long by 12 cm. in circumference), which has been slightly squeezed towards the upper end, bears spirally arranged imperfectly preserved leaf-scars and the lower end shows the truncated base of one of the short Stigmaria-like arms characteristic of the plant. As shown clearly in a specimen originally figured by Bischof and more recently by Potonié[161], the stem-base is divided by a double dichotomy into four short and broad lobes with blunt apices and bent upwards like the arms of a grappling iron ([fig. 134], D). The surface of this basal region is characterised by numerous circular scars ([fig. 134], D; 4 scars enlarged) in the form of slightly projecting areas with a depression in the centre of each. These are undoubtedly the scars of rootlets, remains of which are occasionally seen radiating through the surrounding rock. As seen in [fig. 134], D, a, the fractured surface of a basal area may reveal the existence of an axial vascular cylinder giving off slender branches to the rootlets.

Fig. 134. Pleuromeia Sternbergi.

• A. Cast of stem in the Breslau Museum (⅓ nat. size). (A.C.S.)
• B. “Sigillaria oculina” Blanckenhorn. (After Weiss).
• C, D. Leaf-scars and base of stem: a, vascular tissue. (After Solms-Laubach.)

The bulbous enlargement at the base of the Brown seaweed Laminaria bulbosa Lam.[162] simulates the swollen base of Pleuromeia; but a confusion between these two plants is hardly likely to occur. Above the Stigmaria-like base the gradually tapered axis, in the less decorticated specimens, bears spirally disposed transversely elongated areas consisting of two triangular scars between which is the point of exit of a leaf-trace. The form of the leaf-scars is best seen on the face of a mould figured by Solms-Laubach ([fig. 134], C): in this case the two triangular areas appear as slight projections separated by a narrow groove marking the position of the vascular bundle of the leaf. The curved lines above and below the leaf-scar probably mark the boundary of the leaf-base. The two triangular scars are compared by Solms-Laubach and by Potonié with the parichnos-scars of Sigillaria and Lepidodendron (cf. [fig. 146], C), but the large size of the Pleuromeia scars constitutes an obvious difference though possibly not a distinction of importance.

The occurrence of a vertical canal filled with carbonaceous material in some of the stems throws light on the internal structure: the canal, which is described by Solms-Laubach as having a stellate outline in transverse section recalls the narrow central cylinder of a Lepidodendron stem, and this comparison is strengthened by the presence of obliquely ascending grooves which represent leaf-traces passing through the cortex. In specimens which have lost more of the cortical tissues the surface is characterised by spirally disposed, discontinuous vertical grooves representing portions of leaf-traces precisely as they appear in similar casts of Lepidodendron. There is no direct evidence of the existence of secondary wood in the stem, but, as Potonié has pointed out, the greater transverse elongation of the leaf-scars in the lower part of a cast ([fig. 134], A) points to the production of some secondary tissue either in the vascular cylinder or cortex, or possibly in both regions.

In some specimens of Pleuromeia the upper portion is clothed with crowded and imbricate sporophylls which reach a length of 2·5 cm., a maximum breadth of 2·7 cm., and a thickness of 1 mm. Each sporophyll has a thin wing-like border, and on the lower face are several parallel lines. Solms-Laubach describes the sporangium or ovule as attached to the lower surface of the sporophyll and this opinion has been confirmed by Fitting[163] who has also brought forward satisfactory evidence in favour of the sporangial nature of the reproductive organs. Fitting found numerous spores in the Bunter Sandstone near Halle; these are flattened circular bodies 0·5–0·7 mm. in diameter with a granulated surface and the three converging lines characteristic of spores produced in tetrads. The comparison made by this author between the sporophylls of Pleuromeia, which bore the sporangia on the lower surface instead of on the upper as in other lycopodiaceous plants, and the pollen-sacs of Conifers, is worthy of note in reference to the possible relationship between Conifers and Lycopods.

A comparison of the Isoetes stem represented in [fig. 132], A, with the base of a Pleuromeia shows a striking similarity, but, as Fitting points out, the Stigmaria-like arms of the fossil contained a vascular cylinder whereas the blunt lobes of Isoetes consist exclusively of cortical tissue, the roots being given off from the grooves between the lobes of the tuberous stem.

The position of Pleuromeia must for the present be left an open question; it is, however, clear that the plant bears a close resemblance in the form of its base to the Stigmarian branches of Lepidodendron and Sigillaria. The vegetative shoot appears to be constructed on a plan similar to that of these two Palaeozoic genera, but the strobilus is of a different type. It would seem probable that Pleuromeia may be closely allied to Isoetes and to the arborescent Lycopods of Palaeozoic floras. It is not improbably a link in a chain of types which includes Sigillaria on the one hand and Isoetes on the other.

It is not improbable that a specimen from the Lower Bunter of Commern which Blanckenhorn made the type of a new species, Sigillaria oculina ([fig. 134], B) is specifically identical with Pleuromeia Sternbergi. An examination of a cast of the type-specimen in the Berlin Bergakademie led me to regard the fossil with some hesitation as a true Sigillaria, but a more extended knowledge of Pleuromeia lends support to the view adopted by Potonié[164] that Blanckenhorn’s plant is not genetically distinct from Pleuromeia Sternbergi. The resemblance between Sigillaria oculina and some of the Palaeozoic species of Sigillaria emphasised by Weiss[165] has given rise to the belief that the genus Sigillaria persisted into the Triassic era; it is, however, highly probable that the Bunter specimen has no claim to the generic name under which it has hither to been known.

The Bunter Sandstone in which Pleuromeia is the sole representative of plant-life, at least in certain localities, is usually considered to be a desert formation. We may not be far wrong in accepting Fitting’s suggestion that in this isolated species we have a relic of the sparse vegetation which was able to exist where the presence of lakes added a touch of life to the deadness of the Triassic desert.

Pleuromeia is recorded by Fliche as a rare fossil in the Middle Trias of France in the neighbourhood of Lunéville[166].

Herbaceous fossil species of Lycopodiales.

The history of our knowledge of fossil representatives of the Lycopodiales, as also of the Equisetales, affords a striking illustration of the danger of attempting to found a classification on such differences as are expressed by the terms herbaceous and arborescent in the sense in which they are usually employed. As we have seen[167], the presence of secondary wood in stems of the Palaeozoic plant now known as Calamites led so competent a botanist as Adolphe Brongniart to recognise a distinct generic type Calamodendron, which he placed in the Gymnosperms, reserving the designation Calamities for species in which no indication of secondary thickening had been found.

Similarly, the genus Sigillaria was regarded as a Gymnosperm because it was believed to be distinguished from Lepidodendron by the power of forming secondary vascular tissues; the latter genus, originally thought to be always herbaceous, was classed with the Pteridophytes. At the time when this unnatural separation was made between stems with secondary wood and those in which no secondary wood was known to exist, botanists were not aware of the occurrence of any recent Pteridophyte which shared with the higher plants the power of secondary growth in thickness provided by means of a meristematic zone. It is true that the presence or absence of a cambium does not in practice always coincide with the division into herbaceous and arborescent plants: no one would speak of a Date-Palm as a herbaceous plant despite the absence of secondary wood.

The danger which should be borne in mind, in adopting as a matter of convenience the term herbaceous as a sectional heading, is that it should not be taken to imply a complete inability of the so-called herbaceous types to make secondary additions to their conducting tissues. The specimens on which the species of Lycopodites and Selaginellites, (genera which may be designated herbaceous,) are founded are preserved as impressions and not as petrifications; we can, therefore, base definitions only on habit and on such features as are shown by fertile leaves and sporangia. We are fully justified in concluding from evidence adduced by Goldenberg more than fifty years ago and from similar evidence brought to light by more recent researches, that there existed in the Palaeozoic era lycopodiaceous species in close agreement in their herbaceous habit with the lycopods of present-day floras. It has been suggested[168] that the direct ancestors of the genera Lycopodium and Selaginella are represented by the species of Lycopodites and Selaginellites rather than by Lepidodendron and Sigillaria, the arborescent habit of which has been rendered familiar by the numerous attempts to furnish pictorial reproductions of a Palaeozoic forest. Until we are able to subject the species classed as herbaceous to microscopical examination we cannot make any positive statement as to the correctness of this view, but such facts as we possess lead us to regard the suggestion as resting on a sound basis.

Palaeobotanical literature abounds in records of species of Lycopodites, Lycopodium, Selaginella and Selaginites, which have been so named in the belief that their vegetative shoots bear a greater resemblance to those of recent lycopodiaceous plants than to the foliage shoots of Lepidodendron. Many of these records are valueless: Lepidodendra, twigs of Bothrodendron[169] species of conifers, fern rhizomes, and Aphlebiae[170] have masqueraded as herbaceous lycopods. It is obvious that an attempt to identify fossils presenting a general agreement in habit and leaf-form with recent species of lycopods must be attended with considerable risk of error. Recent Conifers include several species the smaller branches of which simulate the leafy shoots of certain species of Lycopodium and Selaginella, and it is not surprising to find that this similarity has been responsible for many false determinations. Among Mosses and the larger foliose Liverworts there are species which in the condition of imperfectly preserved impressions, might easily be mistaken for lycopodiaceous shoots: an equally close resemblance is apparent in the case of some flowering plants, such as New Zealand species of Veronica, Tafalla graveolens (a Composite), Lavoisiera lycopodiodes Gard.[171] (a species of Melastomaceae), all of which have the habit of Cupressineae among the conifers as well as of certain lycopodiaceous plants. It may be impossible to decide whether fossil impressions of branches, which are presumably lycopodiaceous, bear two kinds of leaves[172] like the great majority of recent species of Selaginella. Selaginella grandis, if seen from the under surface, would appear to have two rows of leaves only and might be confused with a small twig of such a conifer as Dacrydium Kirkii, a New Zealand species.

The New Zealand conifers Dacrydium cupressinum Soland. and Podocarpus dacrydioides Rich. closely simulate species of Selaginellites and Lycopodites: in the British Museum a specimen of the latter species bears a label describing it as Lycopodium arboreum (Sir Joseph Hooker and Dr Solander; 1769). The twigs of the Tasmanian conifer Microcachyrs tetragona Hook. f. are very similar in habit to shoots of the recent Lycopodium tetragonum ([fig. 121], C).

In the description of examples of Lycopodites and Selaginellites I have confined myself to such as appear to be above suspicion either because of the presence of spore-bearing organs or, in a few cases, because the specimens of sterile shoots are sufficiently large to show the form of branching in addition to the texture of the leaves. The two generic names Lycopodites and Selaginellites are employed for fossil species which there are substantial grounds for regarding as representatives of Lycopodium and Selaginella. The designation Selaginellites is adopted only for species which afford evidence of heterospory; the name Lycopodites, on the other hand, is used in a comprehensive sense to include all forms—whether homophyllous or heterophyllous—which are not known to be heterosporous. This restricted use of the generic name Selaginellites is advocated by Zeiller[173], who instituted the genus, and by Halle[174] in his recent paper on herbaceous lycopods.

Lycopodites.

The generic term Lycopodites was used by Brongniart in 1822[175] in describing some Tertiary examples of slender axes clothed with small scale-like leaves which he named Lycopodites squamatus. These are fragments of coniferous shoots. In the Prodrome d’une histoire des végétaux fossiles[176] Brongniart included several Palaeozoic and Jurassic species in Lycopodites and instituted a new genus Selaginites, expressing a doubt as to the wisdom of attempting to draw a generic distinction between the two sets of species. In a later work[177] he recognised only one undoubted species, Lycopodites falcatus. The first satisfactory account of fossils referred to Lycopodites is by Goldenberg[178] who gave the following definition of the genus:—“Branches with leaves spirally disposed or in whorls. Sporangia in the axil of foliage leaves or borne in terminal strobili.”

It was suggested by Lesquereux[179] that Goldenberg’s definition, which was intended to apply to herbaceous species, should be extended so as to include forms with woody stems but which do not in all respects agree with Lepidodendron. Kidston[180] subsequently adopted Lesquereux’s modification of Goldenberg’s definition. We cannot draw any well-defined line between impressions of herbaceous forms and those of small arborescent species. We use the name Lycopodites for such plants as appear to agree in habit with recent species of Lycopodium and Selaginella and which, so far as we know, were not heterosporous: it is highly probable that some of the species so named had the power of producing secondary wood, a power possessed by some recent Pteridophytes which never attain the dimensions of arborescent plants.

It has been shown by Halle[181], who has re-examined several of Goldenberg’s specimens which have been acquired by the Stockholm Palaeobotanical Museum, that some of his species of Lycopodites are heterosporous and therefore referable to Zeiller’s genus Selaginellites.

In 1869 Renault described two species of supposed Palaeozoic Lycopods as Lycopodium punctatum and L. Renaultii[182], the latter name having been suggested by Brongniart to whom specimens were submitted. These species were afterwards recognised by their author as wrongly named and were transferred to the genus Heterangium[183], a determination which is probably correct; it is at least certain that the use of the name Lycopodium cannot be upheld.

We have unfortunately to rely on specimens without petrified tissues for our information in regard to the history of Lycopodites and Selaginellites. Among the older fossils referred to Lycopodites are specimens from Lower Carboniferous rocks at Shap in Westmoreland which Kidston originally described as Lycopodites Vanuxemi[184], identifying them with Goeppert’s Sigillaria Vanuxemi[185] founded on German material. In a later paper Kidston transferred the British specimens of vegetative shoots to a new genus Archaeosigillaria[186].

Lycopodites Stockii Kidston[187].

The plant so named was discovered in Lower Carboniferous strata of Eskdale, Dumfries, Scotland; it is represented by imperfectly preserved shoots bearing a terminal strobilus and was originally described by Kidston as apparently possessing two kinds of foliage leaves borne in whorls. The larger leaves have an ovate cordate lamina with an acuminate apex, while the smaller leaves, which are less distinct, are transversely elongated, and simulate sporangia in appearance. Dr Kidston’s figure of this species has recently been reproduced by Professor Bower[188] who speaks of the supposed smaller leaves as sporangia, a view with which the author of the species agrees. It would appear that this identification is, however, based solely on external resemblance and has not been confirmed by the discovery of any spores. Assuming the sporangial nature of these structures, this Palaeozoic type represents, as Bower points out, a condition similar to that in some recent species of Lycopodium in which sporangia are not confined to a terminal strobilus but occur also in association with ordinary foliage leaves. The strobilus consists of crowded sporophylls which are too imperfect to afford any definite evidence as to their homosporous or heterosporous nature. As Solms-Laubach[189] points out, this type recalls Lycopodium Phlegmaria among recent species.

Lycopodites Reidii Penhallow.

Professor Penhallow[190] instituted this name for a specimen measuring 8 cm. long by 6 mm. in breadth, collected by Mr Reid from the Old Red Sandstone of Caithness, consisting of an axis bearing narrow lanceolate leaves some of which bear sporangia at the base.

Lycopodites Gutbieri Goeppert[191].

1894, Lycopodites elongatus Kidston[192] (not Goldenberg).

The species, figured by Geinitz as Lycopodites Gutbieri[193], from the Coal-Measures of Saxony is probably a true representative of the genus. The Saxon specimens are heterophyllous; the larger lanceolate and slightly falcate leaves arranged in two rows, are 4–5 mm. long while the smaller leaves are one half or one third this size; some of the dichotomously branched shoots terminate in long and narrow strobili not unlike those of Zeiller’s species Selaginellites Suissei[194]. Kidston[195] has included under this specific name some fragments collected by Hemingway from the Upper Coal-Measures of Radstock, Somersetshire, but as only one form of leaf is seen the reasons for adopting Goeppert’s designation are perhaps hardly adequate.

Lycopodites ciliatus Kidston[196].

Under this name Kidston describes a small specimen, obtained by Hemingway from the Middle Coal-Measures of Barnsley in Yorkshire, consisting of a slender forked axis bearing oval-acuminate leaves approximately 5 mm. long with a finely ciliate margin. Associated with the leaves were found spores which Kidston regards as megaspores.

Lycopodites macrophyllus Goldenberg[197].

This species, originally described by Goldenberg from the Coal-Measures of Saarbrücken has been re-examined by Halle[198] who is unable to confirm Goldenberg’s statement as to heterophylly. The shoots closely resemble Selaginellites primaevus[199] (Gold).

Fig. 135. Selaginellites and Lycopodites. (After Halle.)

1. Selaginellites primaevus (Gold.). × 10.
2. Megaspore of Selaginellites elongatus (Gold.). × 50.
3. Lycopodites Zeilleri Halle. (Nat. size.)
4. Selaginellites elongatus (Gold.). × 2.

Lycopodites Zeilleri Halle[200]. Fig. 135, C.

Halle has founded this species on specimens, from the Coal-Measures of Zwickau in Saxony, characterised by dimorphic lanceolate leaves in four rows, the larger being 4–6 mm. long: the smaller leaves have a ciliate edge. A comparison is made with the recent species Selaginella arabica Baker, S. revoluta Bak., and S. armata Bak. in which the leaves are described as ciliate. In the absence of sporangia and spores the species is placed in the genus Lycopodites.

Lycopodites lanceolatus (Brodie). Fig. 136.

• 1845 Naiadita lanceolata, Brodie[201].
  Naiadea acuminata, Buckman[202].
• 1850 Naiadea lanceolata, Buckman[203].
  Naiadea petiolata, Buckman[204].
• 1900 Naiadites acuminatus, Wickes[205].
• 1901 Naiadita lanceolata, Sollas[206] (figures showing habit of the plant).
• 1904 Lycopodites lanceolatus, Seward[207] (figure showing habit of the
  plant).

Fig. 136. Lycopodites lanceolatus (Brodie). (After Miss Sollas. × 40.)
a, Sporangium wall; b, leaf.
c, remains of tubular elements in stem.

Specimens referred to this species were originally recorded by Brodie from Rhaetic rocks in the Severn valley, the name Naiadita being chosen as the result of Lindley’s comparison of the small and delicate leaves with those of recent species of the Monocotyledonous family Naiadaceae. The species may be described as follows:

Plant slender and moss-like in habit. The axis, which is delicate and thread-like, bears numerous linear acuminate or narrow ovate leaves reaching a length of approximately 5 mm. Under a low magnifying power the thin lamina of the leaves is seen to have a superficial layer of polygonal or rectangular cells arranged in parallel series ([fig. 136] b). There is no trace of midrib or stomata. The sporangia are more or less spherical and short-stalked, situated at the base of the foliage leaves and containing numerous tetrads of spores. The spores have a diameter of 0·08 mm.

Buckman founded additional species on differences in the shape of the leaves but, as Miss Sollas has pointed out, such differences as he noticed may be detected on the same axis. It was stated in an earlier chapter[208] that Starkie Gardner, on insufficient evidence, proposed to place Brodie’s plant among the Mosses. The discovery by Mr Wickes of new material at Pylle hill near Bristol afforded an opportunity for a re-examination of the species: this was successfully undertaken by Miss Sollas who was able to dissolve out spores from the matrix by dilute hydrochloric acid, and to recognise the remains of internal structure in the slender axes by exposing successive surfaces with the aid of a hone. It was found that sporangia occurred at the base of some of the leaves containing numerous tetrads of spores, the individual spores having a diameter of 0·08 mm., apparently twice as large as those of any recent species of Lycopodium. Fig. 136 shows a sporangium, a, at the base of a leaf, b. Indications of tubular elements were recognised in the stem and it is noteworthy that although the outlines of epidermal cells on the leaves are well preserved no stomata were found. The leaves of the recent American species Lycopodium alopecuroides Linn. var. aquaticum Spring[209], which lives under water, possess stomata. It is probable that in Lycopodites lanceolatus the leaves had a very thin lamina and may have been similar in structure to those of recent Mosses; the plant possibly lived in very humid situations or grew submerged. Miss Sollas’s investigations afford a satisfactory demonstration of the lycopodiaceous nature of this small Rhaetic species: as I have elsewhere suggested[210], the generic name Lycopodites should be substituted for that of Naiadita. Examples of this species may be seen in the British Museum.

The Rhaetic species from Scania, Lycopodites scanicus Nath.[211] (in litt.), recently re-described by Halle and originally referred by Nathorst to Gleichenia affords another example of the occurrence of a small herbaceous lycopod of Rhaetic age.

Fig. 137. Lycopodites falcatus L. and H. From the Inferior Oolite of Yorkshire. (Nat. size. M.S.)

Lycopodites falcatus Lind. and Hutt. Fig. 137.

• 1831 Lycopodites falcatus, Lindley and Hutton[212].
• 1838 Muscites falcatus, Sternberg[213].
• 1870 Lycopodium falcatum, Schimper[214].

In 1822 Young and Bird[215] figured a specimen from the Inferior Oolite rocks of the Yorkshire coast bearing “small round crowded leaves,” which was afterwards described by Lindley from additional material obtained from Cloughton near Scarborough as Lycopodites falcatus. The example represented in [fig. 137] shows the dichotomously branched shoots bearing two rows of broadly falcate leaves. A careful examination of the type-specimen[216] revealed traces of what appeared to be smaller leaves, but there is no satisfactory proof of heterophylly. No sporangia or spores have been found. This British species has been recorded from Lower Jurassic or Rhaetic rocks of Bornholm[217] and a similar though probably not identical type, Lycopodites Victoriae[218], has been recognised in Jurassic strata of Australia (South Gippsland, Victoria). An Indian plant described by Oldham and Morris[219] from the Jurassic flora of the Rajmahal hills as Araucarites (?) gracilis and subsequently transferred by Feistmantel to Schimper’s genus Cheirolepis[220] may be identical with the Yorkshire species. The Jurassic fragments described by Heer from Siberia as Lycopodites tenerrimus[221] may be lycopodiaceous, but they are of no botanical interest.

Other examples of Mesozoic Lycopods have been recorded, but in the absence of well-preserved shoots and sporangia they are noteworthy only as pointing to a wide distribution of Lycopodites in Jurassic and Cretaceous floras[222].

From Tertiary strata species of supposed herbaceous lycopods have been figured by several authors, one of the best of which is Selaginella Berthoudi Lesq.[223] from Tertiary beds in Colorado. This species agrees very closely in the two forms of leaf with Selaginella grandis, but as the specimens are sterile we have not sufficient justification for the employment of the generic name Selaginellites.

Selaginellites.

This generic name has been instituted by Zeiller[224] for specimens from the coal basis of Blanzy (France). It is applied to heterosporous species with the habit of Selaginella: Zeiller preferred the designation Selaginellites to Selaginella on the ground that the type species differs from recent forms in having more than four megaspores in each megasporangium. It is, however, convenient to extend the term to all heterosporous fossil species irrespective of the spore-output.

Selaginellites Suissei Zeiller.

This species was described in Zeiller’s preliminary note[225] as Lycopodites Suissei, but he afterwards transferred it to the genus Selaginellites. In habit the plant bears a close resemblance to Lycopodites macrophyllus of Goldenberg; the shoots, 1–3 mm. thick, are branched in a more or less dichotomous fashion and bear tetrastichous leaves. The larger leaves reach a length of 4–6 mm. and a breadth of 2–3 mm.; the smaller leaves are described as almost invisible, closely applied to the axis, oval-lanceolate and 1–2 mm. long with a breadth of 0·5–0·75 mm. Long and narrow strobili (15 cm. by 8–10 mm.) terminate the fertile branches; these bear crowded sporophylls with a triangular lamina and finely denticulate margin. Oval sporangia were found on the lower sporophylls containing 16–24 spherical megaspores 0·6–0·65 mm. in diameter. The outer membrane of the spore is characterised by fine anastomosing ridges and thin plates radiating from the apex and forming an equatorial collarette. The microspores have a diameter of 40–60μ and the same type of outer membrane as in the megaspores. The megaspores of the recent species Selaginella caulescens, as figured by Bennie and Kidston[226], resemble those of the Palaeozoic type in the presence of an equatorial flange. It is interesting to find that, in spite of the occurrence of 16–24 megaspores in a single sporangium the size of the fossil spores exceeds that of the recent species.