Ancient Plants Being a Simple Account of the past Vegetation of the Earth and of the Recent Important Discoveries Made in This Realm of Nature

Ancient Plants

[Illustration: Photo. of the specimen in Manchester Museum. THE STUMP OF A _LEPIDODENDRON_ FROM THE COAL MEASURES]

Ancient Plants

BEING A SIMPLE ACCOUNT OF THE PAST VEGETATION OF THE EARTH AND OF THE RECENT IMPORTANT DISCOVERIES MADE IN THIS REALM OF NATURE STUDY

BY MARIE C. STOPES, D.Sc., Ph.D., F.L.S. Lecturer in Fossil Botany, Manchester University Author of “The Study of Plant Life for Young People”

LONDON BLACKIE & SON, Limited, 50 OLD BAILEY, E.C. GLASGOW AND BOMBAY 1910

Preface

The number and the importance of the discoveries which have been made in the course of the last five or six years in the realm of Fossil Botany have largely altered the aspect of the subject and greatly widened its horizon. Until comparatively recent times the rather narrow outlook and the technical difficulties of the study made it one which could only be appreciated by specialists. This has been gradually changed, owing to the detailed anatomical work which it was found possible to do on the carboniferous plants, and which proved to be of great botanical importance. About ten years ago textbooks in English were written, and the subject was included in the work of the honours students of Botany at the Universities. To-day the important bearing of the results of this branch of Science on several others, as well as its intrinsic value, is so much greater, that anyone who is at all acquainted with general science, and more particularly with Botany and Geology, must find much to interest him in it.

There is no book in the English language which places this really attractive subject before the non-specialist, and to do so is the aim of the present volume. The two excellent English books which we possess, viz. Seward’s _Fossil Plants_ (of which the first volume only has appeared, and that ten years ago) and Scott’s _Studies in Fossil Botany_, are ideal for advanced University students. But they are written for students who are supposed to have a previous knowledge of technical botany, and prove very hard or impossible reading for those who are merely acquainted with Science in a general way, or for less advanced students.

The inclusion of fossil types in the South Kensington syllabus for Botany indicates the increasing importance attached to palæobotany, and as vital facts about several of those types are not to be found in a simply written book, the students preparing for the examination must find some difficulty in getting their information. Furthermore, Scott’s book, the only up-to-date one, does not give a complete survey of the subject, but just selects the more important families to describe in detail.

Hence the present book was attempted for the double purpose of presenting the most interesting discoveries and general conclusions of recent years, and bringing together the subject as a whole.

The mass of information which has been collected about fossil plants is now enormous, and the greatest difficulty in writing this little book has been the necessity of eliminating much that is of great interest. The author awaits with fear and trembling the criticisms of specialists, who will probably find that many things considered by them as particularly interesting or essential have been left out. It is hoped that they will bear in mind the scope and aim of the book. I try to present only the structure raised on the foundation of the accumulated details of specialists’ work, and not to demonstrate brick by brick the exposed foundation.

Though the book is not written specially for them, it is probable that University students may find it useful as a general survey of the whole subject, for there is much in it that can only be learned otherwise by reference to innumerable original monographs.

In writing this book all possible sources of information have been consulted, and though Scott’s _Studies_[1] naturally formed the foundation of some of the chapters on Pteridophytes, the authorities for all the general part and the recent discoveries are the numerous memoirs published by many different learned societies here and abroad.

As these pages are primarily for the use of those who have no very technical preliminary training, the simplest language possible which is consistent with a concise style has always been adopted. The necessary technical terms are either explained in the context or in the glossary at the end of the book. The list of the more important authorities makes no pretence of including all the references that might be consulted with advantage, but merely indicates the more important volumes and papers which anyone should read who wishes to follow up the subject.

All the illustrations are made for the book itself, and I am much obliged to Mr. D. M. S. Watson, B.Sc., for the microphotos of plant anatomy which adorn its pages. The figures and diagram are my own work.

This book is dedicated to college students, to the senior pupils of good schools where the subject is beginning to find a place in the higher courses of Botany, but especially to all those who take an interest in plant evolution because it forms a thread in the web of life whose design they wish to trace.

M. C. Stopes.

_December, 1909._

Contents

Chap. Page I. Introductory 1 II. Various Kinds of Fossil Plants 6 III. Coal, the most Important of Plant Remains 22 IV. The Seven Ages of Plant Life 33 V. Stages in Plant Evolution 43 VI. Minute Structure of Fossil Plants— Likenesses to Living Ones 53 VII. ″ ″ Differences from Living Ones 69 VIII. Past Histories of Plant Families— (i) Flowering Plants 79 IX. ″ ″ (ii) Higher Gymnosperms 86 X. ″ ″ (iii) Bennettitales 102 XI. ″ ″ (iv) The Cycads 109 XII. ″ ″ (v) Pteridosperms 114 XIII. ″ ″ (vi) The Ferns 124 XIV. ″ ″ (vii) The Lycopods 133 XV. ″ ″ (viii) The Horsetails 145 XVI. ″ ″ (ix) Sphenophyllales 153 XVII. ″ ″ (x) The Lower Plants 161 XVIII. Fossil Plants as Records of Ancient Countries 168 XIX. Conclusion 174

APPENDIX I. List of Requirements for a Collecting Expedition 183 II. Treatment of Specimens 184 III. Literature 186 Glossary 188 Footnotes 193 Index 193

Ancient Plants

Chapter I Introductory

The lore of the plants which have successively clothed this ancient earth during the thousands of centuries before men appeared is generally ignored or tossed on one side with a contemptuous comment on the dullness and “dryness” of fossil botany.

It is true that all that remains of the once luxuriant vegetation are fragments preserved in stone, fragments which often show little of beauty or value to the untrained eye; but nevertheless these fragments can tell a story of great interest when once we have the clue to their meaning.

The plants which lived when the world was young were not the same as those which live to-day, yet they filled much the same place in the economy of nature, and were as vitally important to the animals then depending on them as are the plants which are now indispensable to man. To-day the life of the modern plants interests many people, and even philosophers have examined the structure of their bodies and have pondered over the great unanswered questions of the cause and the course of their evolution. But all the plants which are now alive are the descendants of those which lived a few years ago, and those again came down through generation after generation from the plants which inhabited the world before the races of men existed. If, therefore, we wish to know and understand the vegetation living to-day we must look into the past histories of the families of plants, and there is no way to do this at once so simple and so direct (in theory) as to examine the remains of the plants which actually lived in that past. Yet when we come to do this practically we encounter many difficulties, which have discouraged all but enthusiasts from attempting the study hitherto, but which in reality need not dismay us.

When Lindley and Hutton, in 1831, began to publish their classical book _The Fossil Flora of Great Britain_, they could give but isolated fragments of information concerning the fossils they described, and the results of their work threw but little light on the theoretical problems of morphology and classification of living plants. Since then great advance has been made, and now the sum of our knowledge of the subject, though far from complete, is so considerable and has such a far-reaching influence that it is becoming the chief inspiration of several branches of modern botany. Of the many workers who have contributed to this stock of knowledge the foremost, as he was the pioneer in the investigations on modern lines, is Williamson, who was a professor at Manchester University, and whose monographs and specimens are classics to-day. Still living is Dr. Scott, whose greatness is scarcely less, as well as an ever-increasing number of specialists in this country, who are continually making discoveries. Abroad, the chief Continental names are Renault, Bertrand, Count Solms Laubach, Brongniart, Zeiller; and in America is Dr. Wieland; while there are innumerable other workers in the field who have deepened and widened the channels of information. The literature on fossil plants is now vast; so great that to give merely the names of the publications would fill a very large volume.

But, like the records left by the plants themselves, most of this literature is unreadable by any but specialists, and its really vital interest is enclosed in a petrifying medium of technicalities. It is to give their results in a more accessible form that the present volume has been written.

The actual plants that lived and died long ago have left either no trace of their form and character, or but imperfect fragments of some of their parts embedded in hard rock and often hidden deep in the earth. That such difficulties lie in our way should not discourage us from attempting to learn all the fossils can teach. Many an old manuscript which is torn and partly destroyed bears a record, the fragments of which are more interesting and important than a tale told by a complete new book. The very difficulty of the subject of fossil botany is in itself an incentive to study, and the obstacles to be surmounted before a view of the ancient plants can be seen increase the fascination of the journey.

The world of to-day has been nearly explored; but the world, or rather the innumerable world-phases of the past, lie before us practically unknown, bewilderingly enticing in their mystery. These untrodden regions are revealed to us only by the fossils lying scattered through the rocks at our feet, which give us the clues to guide us along an adventurous path.

Fables of flying dragons and wondrous sea monsters have been shown by the students of animal fossils to be no more marvellous than were the actual creatures which once inhabited the globe; and among the plants such wonderful monsters have their parallels in the floras of the past. The trees which are living to-day are very recent in comparison with the ancestors of the families of lowlier plants, and most of the modern forest trees have usurped a position which once belonged to the monster members of such families as the Lycopods and Equisetums, which are now humble and dwindling. An ancient giant of the past is seen in the frontispiece, and the great girth of its stem offers a striking contrast to the feeble trailing branches of its living relatives, the Club-mosses.

As we follow their histories we shall see how family after family has risen to dominate the forest, and has in its turn given place to a succeeding group. Some of the families that flourished long since have living descendants of dwarfed and puny growth, others have died out completely, so that their very existence would have been unsuspected had it not been revealed by their broken fragments entombed in the rocks.

From the study of the fossils, also, we can discover something of the course of the evolution of the different parts of the plant body, from the changes it has passed through in the countless ages of its existence. Just as the dominant animals of the past had bodies lacking in many of the characters which are most important to the living animals, so did the early plants differ from those around us to-day. It is the comparative study of living and fossil structures which throws the strongest light on the facts and factors of evolution.

When the study of fossil organisms goes into minute detail and embraces the fine subtleties of their internal structure, then the student of fossil plants has the advantage of the zoological observer, for in many of the fossil plants the cells themselves are petrified with a perfection that no fossil animal tissues have yet been found to approach. Under the microscope the most delicate of plant cells, the patterns on their walls, and sometimes even their nuclei can be recognized as clearly as if they were living tissues. The value of this is immense, because the external appearance of leaves and stems is often very deceptive, and only when both external appearance and internal structure are known can a real estimate of the character of the plant be made. In the following chapters a number of photographs taken through the microscope will show some of the cell structure from fossil plants. Such figures as fig. 11 and fig. 96, for example, illustrate the excellence of preservation which is often found in petrified plant tissues. Indeed, the microscope becomes an essential part of the equipment of a fossil botanist; as it is to a student of living plants. But for those who are not intending to specialize on the subject micro-photographs will illustrate sufficient detail, while in most modern museums some excellently preserved specimens are exhibited which show their structure if examined with a magnifying glass.

We recognize to-day the effect the vegetation of a district has on its scenery, even on its more fundamental nature; and we see how the plants keep in close harmony with the lands and waters, the climates and soils of the places they inhabit. So was it in the past. Hence the fossil plants of a district will throw much light on its physical characters during the epoch when they were living, and from their evidence it is possible to build up a picture of the conditions of a region during the epochs of its unwritten history.

From every point of view a student of living plants will find his knowledge and understanding of them greatly increased by a study of the fossils. Not only to the botanist is the subject of value, the geologist is equally concerned with it, though from a slightly different viewpoint, and all students of the past history of the earth will gain from it a wider knowledge of their specialty.

To all observers of life, to all philosophers, the whole history of plants, which only approaches completion when the fossils are studied, and compared or contrasted with living forms, affords a wonderful illustration of the laws of evolution on which are based most of the modern conceptions of life. Even to those whose profession necessitates purely practical lines of thought, fossil botany has something to teach; the study of coal, for instance, comes within its boundaries. While to all who think on the world at all, the story told by the fossil plants is a chapter in the Book of Life which is as well worth reading as any in that mystical volume.

Chapter II Various Kinds Of Fossil Plants

Of the rocks which form the solid earth of to-day, a very large proportion have been built up from the deposits at the bottom of ancient oceans and lakes. The earth is very old, and in the course of its history dry land and sea, mountains and valleys have been formed and again destroyed on the same spot, and it is from the silt at the bottom of an ocean that the hills of the future are built.

The chief key we have to the processes that were in operation in the past is the course of events passing under our eyes to-day. Hence, if we would understand the formation of the rocks in the ancient seas, we must go to the shores of the modern ones and see what is taking place there. One of the most noticeable characters of a shore is the line of flotsam that is left by the edge of the waves; here you may find all kinds of land plants mixed with the sea shells and general rubbish, plants that may have drifted far. Much of the débris (outside towns) is brought down by the rivers, and may be carried some distance out to sea; then part becomes waterlogged and sinks, and part floats in to shore, perhaps to be carried out again, or to be buried under the coarse sand of the beach. When we examine sandstone rock, or the finer grained stones which are hardened mud, we find in them the remains of shells, sometimes of bones, and also of plant leaves and stems, which in their time had formed the flotsam of a shore. Indeed, one may say that nearly every rock which has not been formed in ancient volcanoes, or been altered by their heat, carries in it _some_ trace of plant or animal. These remains are often very fragmentary and difficult to recognize, but sometimes they are wellnigh as perfect as dried specimens of living things. When they are recognizable as plant or animal remains they are commonly called “fossils”, and it is from their testimony that we must learn all we can know about the life of the past.

[Illustration: Fig. 1.—The Face of a Quarry, showing layers or “beds” of different rock, _a_, _b_, and _c_. The top gravel and soil _s_ has been disintegrated by the growing plants and atmosphere.]

If we would find such stones for ourselves, the quarries offer the best hunting ground, for there several layers of rock are exposed, and we can reach fresh surfaces which have not been decayed by rain and storm. Fig. 1 shows a diagram of a quarry, and illustrates the almost universal fact that the beds of rock when undisturbed lie parallel to each other. Rock _a_ in the figure is fine-grained limestone, _b_ black friable shale mixed with sand, and _c_ purer shale. In such a series of rocks the best fossils will be found in the limestone; its harder and finer structure acting as a better preservative of organisms than the others. In limestone one finds both plant and animal fossils, very often mixed together as the flotsam on the shore is mixed. Many limestones split along parallel planes, and may break into quite thin sheets on whose surfaces the flattened fossils show particularly well.

It is, however, with the plant fossils that we must concern ourselves, and among them we find great variety of form. Some are more or less complete, and give an immediate idea of the size and appearance of the plant to which they had belonged; but such are rare. One of the best-known examples of this type is the base of a great tree trunk illustrated in the frontispiece. With such a fossil there is no shadow of doubt that it is part of a giant tree, and its spreading roots running so far horizontally along the ground suggest the picture of a large crown of branches. Most fossils, however, are much less illuminating, and it is usually only by the careful piecing together of fragments that we can obtain a mental picture of a fossil plant.

A fossil such as that illustrated in the frontispiece—and on a smaller scale this type of preservation is one of the commonest—does not actually consist of the plant body itself. Although from the outside it looks as though it were a stem base covered with bark, the whole of the inner portion is composed of fine hard rock with no trace of woody tissue. In such specimens we have the shape, size, and form of the plant preserved, but none of its actual structure or cells. It is, in fact, a Cast. Fossil casts appear to have been formed by fine sand or mud silting round a submerged stump and enclosing it as completely as if it had been set in plaster of Paris; then the wood and soft tissue decayed and the hollow was filled up with more fine silt; gradually all the bark also decayed and the mud hardened into stone. Thus the stone mould round the outside of the plant enclosed a stone casting. When, after lying for ages undisturbed, these fossils are unearthed, they are so hard and “set” that the surrounding stone peels away from the inner part, just as a plaster cast comes away from an object and retains its shape. There are many varieties of casts among fossil plants. Sometimes on breaking a rock it will split so as to show the perfect form of the surface of a stem, while its reverse is left on the stone as is shown in fig. 2. Had we only the reverse we should still have been able to see the form of the leaf bases by taking a wax impression from it; although there is nothing of the actual tissue of the plant in such a fossil. Sometimes casts of leaf bases show the detail preserved with wonderful sharpness, as in fig. 3. This is an illustration of the leaf scars of _Lepidodendron_, which often form particularly good casts.

[Illustration: Fig. 2.—A, Cast of the Surface showing the Shape of Leaf Bases of _Sigillaria_; B, the reverse of the impression left on the adjacent layer of rock. (Photo.)]

In other instances the cast may simply represent the internal hollows of the plant. This happens most commonly in the case of stems which contained soft pith cells which quickly decayed, or with naturally hollow stems like the Horse-tails (_Equisetum_) of to-day. Fine mud or sand silted into such hollows completely filling them up, and then, whether the rest of the plant were preserved or not, the shape of the inside of the stem remains as a solid stone. Where this has happened, and the outer part of the plant has decayed so as to leave no trace, the solid plug of stone from the centre may look very much like an actual stem itself, as it is cylindrical and may have surface markings like those on the outsides of stems. Some of the casts of this type were for long a puzzle to the older fossil botanists, particularly that illustrated in fig. 4, where the whole looks like a pile of discs.

[Illustration: Fig. 3.—Cast of the Leaf Bases of _Lepidodendron_, showing finely marked detail. (Photo.)]

[Illustration: Fig. 4.—“_Sternbergia._” Internal cast of the stem of _Cordaites_.]

The true nature of this fossil was recognized when casts of the plan were found with some of the wood preserved outside the castings; and it was then known that the plant had a hollow pith, with transverse bands of tissue across it at intervals which caused the curious constrictions in the cast.

[Illustration: Fig. 5.—Leaf Impressions of “Fern” _Sphenopteris_ on Shale. (Photo.)]

Another form of cast which is common in some rocks is that of seeds. As a rule these casts are not connected with any actually preserved tissue, but they show the external form, or the form of the stony part of the seed. Well-known seeds of this type are those of _Trigonocarpon_, which has three characteristic ridges down the stone. Sometimes in the fine sandstone in which they occur embedded, the _internal_ cast lies embedded _in the external_ cast, and between them there is a slight space, now empty, but which once contained the actual shell of the seed, now decayed. Thus we may rattle the “stone” of a fossil fruit as we do the dried nuts of to-day—the external resemblance between the living and the fossil is very striking, but of the actual tissues of the fossil seed nothing is left.

Casts have been of great service to the fossil botanists, for they often give clear indications of the external appearance of the parts they represent; particularly of stems, leaf scars, and large seeds. But all such fossils are very imperfect records of the past plants, for none of the actual plant tissues, no minute anatomy or cell structure, is preserved in that way.

A type of fossil which often shows more detail, and which usually retains something of the actual tissues of the plant, is that known technically as the Impression. These fossils are the most attractive of all the many kinds we have scattered through the rocks, for they often show with marvellous perfection the most delicate and beautiful fern leaves, such as in fig. 5. Here the plant shows up as a black silhouette against the grey stone, and the very veins of the midrib and leaves are quite visible.

Fig. 6 shows another fernlike leaf in an impression, not quite flat like that shown in fig. 5, but with a slight natural curvature of the leaves similar to what would have been their form in life. Though an impression, this specimen is not of the “pressed plant” type, it almost might be described as a _bas-relief_.

Sometimes impressions of fern foliage are very large, and show highly branched and complex leaves like those of tree ferns, and they may cover large sheets of stone. They are particularly common in the fine shales above coal seams, and are best seen in the mines, for they are often too big to bring to the surface complete.

In most impressions the black colour is due to a film of carbon which represents the partly decomposed tissues of the plant. Sometimes this film is cohesive enough to be detached from the stone without damage. Beautiful specimens of this kind are to be seen in the Royal Scottish Museum, Edinburgh where the coiled bud of a young fern leaf has been separated from the rock on which it was pressed, and mounted on glass. Such specimens might be called mummy plants, for they are the actual plant material, but so decayed and withered that the internal cells are no longer intact. In really well preserved ones it is sometimes possible to peel off the plant film, and then treat it with strong chemical agents to clear the black carbon atoms away, and mount it for microscopic examination, when the actual outline of the epidermis cells can be seen.

[Illustration: Fig. 6.—Impression of _Neuropteris_ Leaf, showing details of veins, the leaves in partial relief. (Photo.)]

[Illustration: Fig. 7.—Leaf Impression of _Ginkgo_, of which the film was strong enough to peel off complete]

In fig. 7, the impression is that of a _Ginkgo_ leaf, and after treatment the cells of the epidermis were perfectly recognizable under the microscope, with the stomates (breathing pores) also well preserved. This is shown in fig. 8, where the outline of the cells was drawn from the microscope. In such specimens, however, it is only the outer skin which is preserved, the inner soft tissue, the vital anatomy of the plant, is crushed and carbonized.

Leaves, stems, roots, even flowers (in the more recent rocks) and seeds may all be preserved as impressions; and very often those from the more recently formed rocks are so sharply defined and perfect that they seem to be actual dried leaves laid on the stone.

[Illustration: Fig. 8.—Outline of the Cells from Specimen of Leaf shown in fig. 7

_c_, Ordinary cells; _s_, stomates; _v_, elongated cells above the vein.]

Much evidence has been accumulated that goes to show that the rocks which contain the best impressions were originally deposited under tranquil conditions in water. It might have been in a pool or quiet lake with overshadowing trees, or a landlocked inlet of the sea where silt quietly accumulated, and as the plant fragments fell or drifted into the spot they were covered by fine-grained mud without disturbance. In the case of those which are very well preserved this must have taken place with considerable rapidity, so that they were shut away from contact with the air and from the decay which it induces.

Impressions in the thin sheets of fine rock may be compared to dried specimens pressed between sheets of blotting paper; they are flattened, preserved from decay, and their detailed outline is retained. Fossils of this kind are most valuable, for they give a clear picture of the form of the foliage, and when, as sometimes happens, large masses of leaves, or branches with several leaves attached to them, are preserved together, it is possible to reconstruct the plant from them. It is chiefly from such impressions that the inspiration is drawn for those semi-imaginary pictures of the forests of long ago. From them also are drawn many facts of prime importance to scientists about the nature and appearance of plants, of which the internal anatomy is known from other specimens, and also about the connection of various parts with each other.

Sometimes isolated impressions are found in clay balls or nodules. When the latter are split open they may show as a centre or nucleus a leaf or cone, round which the nodule has collected. In such cases the plant is often preserved without compression, and may show something of the minute details of organization. The preservation, however, is generally far from perfect when viewed from a microscopical standpoint. Fig. 9 shows one of these smooth, clayey nodules split open, and within it the cone which formed its centre, also split into two, and standing in high relief, with its scales showing clearly. Similar nodules or balls of clay are found to-day, forming in slowly running water, and it may be generally observed that they collect round some rubbish, shell, or plant fragment. These nodules are particularly well seen nowadays in the mouth of the Clyde, where they are formed with great rapidity.

[Illustration: Fig. 9.—Clay Nodule split open, showing the two halves of the cone which was its centre. (Photo.)]

Another kind of preservation is that which coats over the whole plant surface with mineral matter, which hardens, and thus preserves the _form_ of the plant. This process can be observed going on to-day in the neighbourhood of hot volcanic streams where the water is heavily charged with minerals. In most cases such fossils have proved of little importance to science, though there are some interesting specimens in the French museums which have not yet been fully examined. A noteworthy fossil of this type is the _Chara_, which, growing in masses together, has sometimes been preserved in this way in large quantities, indicating the existence of an ancient pond in the locality.

There is quite a variety of other types of preservation among fossil plants, but they are of minor interest and importance, and hardly justify detailed consideration. One example that should be mentioned is Amber. This is the gum of old resinous trees, and is a well-known substance which may rank as a “fossil”. Jet, too, is formed from plants, while coal is so important that the whole of the next chapter will be devoted to its consideration. Even the black lead of pencils possibly represents plants that were once alive on this globe.

Though such remains tell us of the existence of plants at the place they were found at a known period in the past, yet they tell very little about the actual structure of the plants themselves, and therefore very little that is of real use to the botanist. Fortunately, however, there are fossils which preserve every cell of the plant tissues, each one perfect, distended as in life, and yet replaced by stone so as to be hard and to allow of the preparation of thin sections which can be studied with the microscope. These are the vegetable fossils which are of prime importance to the botanist and the scientific enquirer into the evolution of plants. Such specimens are commonly known as Petrifactions.

Sometimes small isolated stumps of wood or branches have been completely permeated by silica, which replaces the cell walls and completely preserves and hardens the tissues. This silicified wood is found in a number of different beds of rock, and may be seen washed out on the shore in Yorkshire, Sutherland, and other places where such rocks occur. When such a block is cut and polished the annual rings and all the fine structure or “grain” of the wood become as apparent as in recent wood. From these fossils, too, microscopic sections can be cut, and then the individual wood cells can be studied almost as well as those of living trees. A particularly notable example of fossil tree trunks is the Tertiary forest of the Yellowstone Park. Here the petrified trunks are weathered out and stand together much as they must have stood when alive; they are of course bereft of their foliage branches.

Such specimens, however, are usually only isolated blocks of wood, often fragments from large stumps which show nothing but the rings of late-formed wood. It is impossible to connect them with the impressions of leaves or fruits in most cases, so that of the plants they represent we know only the anatomical structure of the secondary wood and nothing of the foliage or general appearance of the plant as a whole. Hence these specimens also give a very partial representation of the plants to which they belonged.

Fortunately, however, there is still another type of preservation of fossils, a type more perfect than any of the others and sometimes combining the advantages of all of them. This is the special type of petrifaction which includes, not a single piece of wood, but a whole mass of vegetation consisting of fragments of stems, roots, leaves, and even seeds, sometimes all together. These petrifactions are those of masses of forest débris which were lying as they dropped from the trees, or had drifted together as such fragments do. The plant tissues in such masses are preserved so that the most delicate soft tissue cells are perfect, and in many cases the sections are so distinct that one might well be deluded into the belief that it is a living plant at which one looks.

Very important and well-known specimens have been found in France and described by the French palæobotanists. As a rule these specimens are preserved in silica, and are found now in irregular masses of the nature of chert. Of still greater importance, however, owing partly to their greater abundance and partly to the quantity of scientific work that has been done on them, are the masses of stone found in the English coal seams and commonly called “coal balls”.

The “coal balls” are best known from Lancashire and Yorkshire, where they are extremely common in some of the mines, but they also occur in Westphalia and other places on the Continent.

[Illustration: Fig. 10.—Mass of Coal with many “coal balls” embedded in it

_a a_, In surface view; _b b_, cut across. All washed with acid to make the coal balls show up against the black coal. (Photo by Lomax.)]

In external appearance the “coal balls” are slightly irregular roundish masses, most generally about the size of potatoes, and black on the outside from films of adhering coal. Their size varies greatly, and they have been found from that of peas up to masses with a diameter of a foot and a half. They lie embedded in the coal and are not very easily recognizable in it at first, because they are black also, but when washed with acid they turn greyish-white and then can be recognized clearly. Fig. 10 shows a block of coal with an exceptionally large number of the “coal balls” embedded in it. This figure illustrates their slightly irregular rounded form in a typical manner. By chemical analysis they are found to consist of a nearly pure mixture of the carbonates of lime and magnesia; though in some specimens there is a considerable quantity of iron sulphide, and in all there is at least 5 per cent of various impurities and some quantity of carbon.

The important mineral compounds, CaCO_3 and MgCO_3, are mixed in very different quantities, and even in coal balls lying quite close to each other there is often much dissimilarity in this respect. In whatever proportion these minerals are combined, it seems to make but little difference to their preservative power, and in good “coal balls” they may completely replace and petrify each individual cell of the plants in them.

[Illustration: Fig. 11.—Photograph of Section across Stem of _Sphenophyllum_ from a Lancashire “coal ball”, showing perfect preservation of woody tissue

W, wood; _c_, cortex.]

Fig. 11 shows a section across the wood of a stem preserved in a “coal ball”, and illustrates a degree of perfection which is not uncommon. In the course of the succeeding chapters constant reference will be made to tissues preserved in “coal balls”, and it may be noticed that not only the relatively hard woody cells are preserved but the very softest and youngest tissues also appear equally unharmed by their long sojourn in the rocks.

[Illustration: Fig. 12.—Photograph of Section through a Bud of _Lepidodendron_, showing many small leaves tightly packed round the axis. From a “coal ball”]

The particular value of the coal balls as records of past vegetation lies in the fact that they are petrifactions, not of individual plants alone, but of masses of plant débris. Hence in one of these stony concretions may lie twigs with leaves attached, bits of stems with their fruits, and fine rootlets growing through the mass. A careful study and comparison of these fragments has led to the connection, piece by piece, of the various parts of many plants. Such a specimen as that figured in fig. 12 shows how the soft tissues of young leaves are preserved, and how their relation to each other and to the axis is indicated.

Hitherto the only concretions of the nature of “coal balls” containing well preserved plant débris, have been found in the coal or immediately above it, and are of Palæozoic age (see p. 34). Recent exploration, however, has resulted in the discovery of similar concretions of Mesozoic age, from which much may be hoped in the future. Still, at present, it is to the palæozoic specimens we must turn for nearly all valuable knowledge about ancient plants, and primarily to that form of preservation of the specimens known as structural petrifactions, of which the “coal balls” are both the commonest and the most perfect examples.

Chapter III Coal, The Most Important Of Plant Remains

Some of the many forms which are taken by fossil plants were shortly described in the last chapter, but the most important of all, namely coal, must now be considered. Of the fossils hitherto mentioned many are difficult to recognize without examining them very closely, and one might say that all have but little influence on human life, for they are of little practical or commercial use, and their scientific value is not yet very widely known. Of all fossil plants, the great exception is coal. Its commercial importance all over the world needs no illustration, and its appearance needs no description for it is in use in nearly every household. Quite apart from its economic importance, coal has a unique place among fossils in the eyes of the scientist, and is of special interest to the palæontologist.

In England nearly all the coal lies in rocks of a great age, belonging to a period very remote in the world’s history. The rocks bearing the coal contain other fossils, principally those of marine animals, which are characteristic of them and of the period during which they were formed, which is generally known as the “Coal Measure period”. There is geological proof that at one time the coal seams were much more widely spread over England than they are at present; they have been broken up and destroyed in the course of ages, by the natural movements among the rocks and by the many changes and processes of disintegration and decay which have gone on ever since they were deposited. To-day there are but relatively small coal-bearing areas, which have been preserved in the hollows of the synclines.[2]

The seams of coal are extremely numerous, and even the same seam may vary greatly in thickness. From a quarter of an inch to five or six feet is the commonest thickness for coal in this country, but there are many beds abroad of very much greater size. Thin seams often lie irregularly in coarse sandstone; for example, they may be commonly seen in the Millstone Grit; but typical coal seams are found embedded between rocks of a more or less definite character known as the “roof” and “floor”.

[Illustration: Fig. 13.—Diagram of a Series of Parallel Coal Seams with Underclays and Shale Roofs of varying thicknesses]

Basalts, granites, and such rocks do not contain coal; the coal measures in which the seams of coal occur are, generally speaking, limestones, fine sandstones, and shales, that is to say, rocks which in their origin were deposited under water. In detail almost every seam has some individual peculiarity, but the following represents two types of typical seams. In many cases, below the coal, the limestone or sandstone rocks give place to fine, yellow-coloured layers of clay, which varies from a few inches to many feet in thickness and is called the “underclay”. This fine clay is generally free from pebbles and coarse débris of all kinds, and is often supposed to be the soil in which the plants forming the coal had been growing. The line of demarcation between the coal and the clay is usually very sharp, and the compact black layers of hard coal stop almost as abruptly on the upper side and give place to a shale or limestone “roof”; see fig. 13, layers 5, 6, and 7. Very frequently a number of small seams come together, lying parallel, and sometimes succeeding each other so rapidly that the “roof” is eliminated, and a clay floor followed by a coal seam, is succeeded immediately by another clay floor and another coal seam, as in fig. 13, layers 10, 11, and 12. The relative thickness of these beds also varies very greatly, and over an underclay of seven or eight feet the coal seam may only reach a couple of inches, while a thick seam may have a floor of very slight dimensions. These relations depend on such a variety of local circumstances from the day they were forming, that it is only possible to unravel the causes when an individual case is closely studied. The main sequence, however, is constant and is that illustrated in fig. 13.

The second type of seam is that in which the underclay floor is not present, and is replaced either by shales or by a special very hard rock of a finely granular nature called “gannister”. In the gannister floor it is usual to find traces of rootlets and basal stumps of plants, which seem to indicate that the gannister was the ground in which the plants forming the coal were rooted. The coal itself is generally very pure plant remains, though between its layers are often found bands of shaly stone which are called “dirt bands”. These are particularly noticeable in thick seams, and they may be looked on as corresponding to the roof shales; as though, in fact, the roof had started to form but had only reached a slight development when the coal formation began again.

[Illustration: Fig. 14.—Diagram of Coal Seam with Gannister Floor, in which are traces of rootlets _r_, and of stumps of root-like organs _s_]

That the coal is strikingly different from the rocks in which it lies is very obvious, but that alone is no indication of its origin. It is now so universally known and accepted that coal is the remains of vegetables that no proofs are usually offered for the statement. It is, however, of both interest and importance to marshal the evidence for this belief. The grounds for recognizing coal as consisting of practically pure plant remains are many and various, so that only the more important of them will be considered now. The most direct suggestion lies in the impressions of leaves and stems which are found between its layers; this, however, is confronted by the parallel case of plant impressions found in shales and limestones which are not of vegetable origin, so that it might be argued that those plants in the coal drifted in as did those in the limestone. But when we examine the black impressions on limestone or sandstone, an item of value is noticeable; it is often possible to peel off a film, lying between the upper and lower impression, of black coaly substance, sometimes an eighth of an inch thick, and hard and shining like coal. This follows the outline of the plant form of the impression, and it is certain that this minute “coal seam” was formed from the plant tissues. It is, in fact, a coal seam bearing the clearest possible evidence of its plant nature. We have only to imagine this multiplied by many plants lying tightly packed together, with no mineral impurities between, to see that it would yield a coal seam like those we find actually existing.

In some cases in the coal itself a certain amount of the structure of the plants which formed it remains, though usually, in the process of their decay the tissues have entirely decomposed, and left only their carbonized elements. Chemical analysis reveals that, beyond the percentage of mineral ash which is found in living plants, there is little in a pure sample of coal that is not carbonaceous. All the deposits of carbon found in any form in nature can be traced to some animal or vegetable remains, so that it is logical to assume that coal also arose from either animal or plant débris. But were coal of an animal origin, the amount of mineral matter in it would be much larger as well as being of a different nature; for almost all animals have skeletons, even the simplest single-celled protozoa often own calcareous shells, sponges have siliceous spicules, molluscs hard shells, and the higher animals bones and teeth. These things are of a very permanent nature, and would certainly be found in quantities in the coal had animals formed it. Further, the peat of to-day, which collects in thick compact masses of vegetable, shows how plants may form a material consisting of carbonized remains. By certain experiments in which peat was subjected to pressure and heat, practically normal coal was made from it.

[Illustration: Fig. 15.—Part of a Coal Ball, showing the concentric bandings in it which are characteristic of concretions]

[Illustration: Fig. 16.—Mass of Coal with Coal Balls, A and B both enclosing part of the same stem L]

Still a further witness may be found in the structure of the “coal balls” described in the last chapter. These stony masses, lying in the pure coal, might well be considered as apart from it and bearing no relation to its structure; but recent work has shown that they were actually formed at the same time as the coal, developing in its mass as mineral concretions round some of the plants in the soft, saturated, peaty mass which was to be hardened into coal later on.[3] All “coal balls” do not show their concretionary structure so clearly, but sometimes it can be seen that they are made with concentric bands or markings like those characteristic of ordinary mineral concretions (see fig. 15). Concretions are formed by the crystallization of minerals round some centre, and it must have happened that in the coal seams in which the coal-ball concretions are found that this process took place in the soft plant mass before it hardened. Recent research has found that there is good evidence that those seams[4] resulted from the slow accumulation of plant débris under the salt or brackish water in whose swamps the plants were growing, and that as they were collecting the ground slowly sank till they were quite below the level of the sea and were covered by marine silt. At the same time some of the minerals present in the sea water, which must have saturated the mass, crystallized partly and deposited themselves round centres in the plant tissues, and by enclosing them and penetrating them preserved them from decay till the mineral structure entirely replaced the cells, molecule by molecule. Evidence is not wanting that this process went on without disturbance, for in fig. 16 is shown a mass of coal in which lie several coal balls, two of which enclose parts of the same plant. This means that round different centres in the same stem two of the concretions were forming and preserving the tissues; the two stone masses, however, did not enlarge enough to unite, but left a part of the tissue unmineralized, which is now seen as a streak of coal. We have here the most important proof that the coal balls are actually formed in the coal and of the plants making the coal, for had those coal balls come in as pebbles, or in any way from the outside into the coal, they could not have remained in such a position as to lie side by side enclosing part of the same stem. There are many other details which may be used in this proof, but this one illustration serves to show the importance of coal balls when dealing with the theories of the origin of coal, for they are perfectly preserved samples of what the whole coal mass was at one time.

There are but few seams, however, which contain coal balls, and about those in which they do not occur our knowledge is very scanty. It is often assumed that the plant impressions in the shales above the coal seams can be taken as fair samples of those which formed the coal itself; but this has been recently shown to be a fallacious argument in some cases, so that it is impossible to rely on it in general. The truth is, that though coal is one of the most studied of all the geological deposits, we are still profoundly ignorant of the details of its formation except in a few cases.

The way in which coal seams were formed has been described often and variously, and for many years there were heated discussions between the upholders of the different views as to the merits of their various theories. It is now certain that there must have been at least four principal ways in which coal was formed, and the different seams are illustrations of the products of different methods. In all cases more or less water is required, for coal is what is known as a sedimentary deposit, that is, one which collects under water, like the fine mud and silt and débris in a lake. It will be understood, however, that if the plant remains were collecting at any spot, and the water brought in sand and mud as well, then the deposit could not have resulted in pure coal, but would have been a sandy mixture with many plant remains, and would have resulted in the formation of a rock, such as parts of the millstone grit, where there are many streaks of coal through the stone.

Among various coal seams, evidence for the following modes of coal formation can be found:—

(_a_) _In fresh water._—In still freshwater lakes or pools, with overhanging plants growing on the banks, twigs and leaves which fell or were blown into the water became waterlogged and sank to the bottom. With a luxuriant growth of plants rapidly collecting under water, and there preserved from contact with the air and its decaying influence, enough plant remains would collect to form a seam. After that some change in the local conditions took place, and other deposits covered the plants and began the accumulations which finally pressed the vegetable mass into coal.

To freshwater lakes of large size plants might also have been brought by rivers and streams; they would have become waterlogged in time, after floating farther than the sand and stones with which they came, and would thus settle and form a deposit practically free from anything but plant remains.

(_b_) _As peat._—Peat commonly forms on our heather moors and bogs to-day to a considerable thickness. This also took place long ago in all probability, and when the level of the land altered it would have been covered by other deposits, pressed, and finally changed into coal.

(_c_) _In salt or brackish water, growing in situ._—Trees and undergrowth growing thickly together in a salt or brackish marsh supplied a large quantity of débris which fell into the mud or water below them, and were thus shut off from the air and partly preserved. When conditions favoured the formation of a coal seam the land level was slowly sinking, and so, though the débris collected in large quantities, it was always kept just beneath the water level. Finally the land sank more rapidly, till the vegetable mass was quite under sea water, then mud was deposited over it, and the materials which were afterwards hardened to form the roof rocks were deposited. This was the case in those seams in which “coal balls” occur, and the evidence of the sea water covering the coal soon after it was deposited lies in the numerous sea shells found in the roof immediately above it.

(_d_) _In salt water, drifted material._—Tree trunks and large tangled masses of vegetation drifted out to sea by the rivers just as they do to-day. These became waterlogged, and finally sank some distance from the shore. (Those sinking near the shore would not form pure coal, for sand and mud would be mixed with them, also brought down by rivers and stirred up from the bottom by waves.) The currents would bring numbers of such plants to the same area until a large mass was deposited on the sea floor. Finally the local conditions would have changed, the currents then bringing mud or sand, which covered the vegetable mass and formed the mineral roof of the resulting coal seam. There is a variety of what might be called the “drifted coals”, which appears to have been formed of nothing but the _spores_ of plants of a resinous nature. These structures must have been very light, and possibly floated a long distance before sinking.

If we could but obtain enough evidence to understand each case fully we should probably find that every coal seam represents some slightly different mode of formation, that in each case there was some local peculiarity in the plants themselves and the way they accumulated in coal-forming masses, but the above four methods will be found to cover the principal ways in which coal has arisen.

Coal, as we now know it, has a great variety of qualities. The differences probably depend only to a small extent on the varieties among the plants forming it, and are almost entirely due to the many later conditions which have affected the coal after its original formation. Some such conditions are the various upheavals and depressions to which the rocks containing the coal have been subjected, the weight of the beds lying over the coal seams, and the high temperatures to which they may have been subjected when lying under a considerable depth of later-deposited rocks. The influence on the coal of these and many other physical factors has been enormous, but they are purely cosmical and belong to the special realm of geological study, and so cannot be considered in detail now.

To return to our special subject, namely, the plants themselves which are now preserved in the coal. Their nature and appearance, their affinities and minute structure, can only be ascertained by a detailed study, to which the following chapters will be devoted, though in their limited space but an outline sketch of the subject can be drawn.

It has been stated by some writers that in the Coal Measure period plants were more numerous and luxuriant than they ever were before or ever have been since. This view could only have been brought forward by one who was considering the geology of England alone, and in any case there appears to be very little real evidence for such a view. Certainly in Europe a large proportion of the coal is of this age, and to supply the enormous masses of vegetation it represents a great growth of plants must have existed. But it is evident that just at the Carboniferous period in what is now called Europe the physical conditions of the land which roughly corresponded to the present Continent were such as favoured the accumulation of plants, and the gradual sinking of the land level also favoured their preservation under rapidly succeeding deposits. Of the countless plants growing in Europe to-day very few stand any chance of being preserved as coal for the future; so that, unless the physical conditions were suitable, plants might have been growing in great quantity at any given period without ever forming coal. But now that the geology of the whole world is becoming better known, it is found that coal is by no means specially confined to the Coal Measure age. Even in Europe coals of a much later date are worked, while abroad, especially in Asia and Australia, the later coals are very important. For example, in Japan, seams of coal 14, 20, and even more feet in thickness are worked which belong to the Tertiary period (see p. 34), while in Manchuria coal 100 feet thick is reported of the same age. When these facts are considered it is soon found that all the statements made about the unique vegetative luxuriance of the Coal Measure period are founded either on insufficient evidence or on no evidence at all.

The plants forming the later coals must have had in their own structure much that differed from those forming the old coals of Britain, and the gradual change in the character of the vegetation in the course of the succeeding ages is a point of first-rate importance and interest which will be considered shortly in the next chapter.

Chapter IV The Seven Ages Of Plant Life

Life has played its important part on the earth for countless series of years, of the length of whose periods no one has any exact knowledge. Many guesses have been made, and many scientific theories have been used to estimate their duration, but they remain inscrutable. When numbers are immense they cease to hold any meaning for us, for the human mind cannot comprehend the significance of vast numbers, of immense space, or of æons of time. Hence when we look back on the history of the world we cannot attempt to give even approximate dates for its events, and the best we can do is to speak only of great periods as units whose relative position and whose relative duration we can estimate to some extent.

Those who have studied geology, which is the science of the world’s history since its beginning, have given names to the great epochs and to their chief subdivisions. With the smaller periods and the subdivisions of the greater ones we will not concern ourselves, for our study of the plants it will suffice if we recognize the main sequence of past time.

The main divisions are practically universal, and evidence of their existence and of the character of the creatures living in them can be found all over the world; the smaller divisions, however, may often be local, or only of value in one continent. To the specialist even the smallest of them is of importance, and is a link in the chain of evidence with which he cannot dispense; but we are at present concerned only with the broad outlines of the history of the plants of these periods, so will not trouble ourselves with unnecessary details.[5] Corresponding to certain marked changes in the character of the vegetation, we find seven important divisions of geological time which we will take as our unit periods, and which are tabulated as follows:—

Cainozoic I. Present Day. II. Tertiary. Mesozoic III. Upper Cretaceous (or Chalk). IV. The rest of the Mesozoic. V. Newer Palæozoic, including Permian. Carboniferous. Devonian. Palæozoic VI. Older Palæozoic. Eozoic VII. Archæan.

Now the actual length of these various periods was very different. The epoch of the Present Day is only in its commencement, and is like a thin line if compared with the broad bands of the past epochs. By far the greatest of the periods is the Archæan, and even the Older Palæozoic is probably longer than all the others taken together. It is, however, so remote, and the rocks which were formed in it retain so little plant structure that is decipherable, so few specimens which are more than mere fragments, that we know very little about it from the point of view of the plant life of the time. It includes the immense indefinite epochs when plants began to evolve, and the later ones when animals of many kinds flourished, and when plants, too, were of great size and importance, though we are ignorant of their structure. Of all the seven divisions of time, we can say least about the two earliest, simply for want of anything to say which is founded on fact rather than on theoretical conclusions.

Although these periods seem clearly marked off from one another when looked at from a great distance, they are, of course, but arbitrary divisions of one long, continuous series of slow changes. It is not in the way of nature to make an abrupt change and suddenly shut off one period—be it a day or an æon—from another, and just as the seasons glide almost imperceptibly into one another, so did the great periods of the past. Thus, though there is a strong and very evident contrast between the plants typical of the Carboniferous period and of the Mesozoic, those of the Permian are to some extent intermediate, and between the beginning of the Permian and the end of the Carboniferous—if judged by the flora—it is often hard to decide.

It must be realized that almost any given spot of land—the north of England, for example—has been beneath the sea, and again elevated into the air, at least more than once. That the hard rocks which make its present-day hills have been built up from the silt and débris under an ocean, and after being formed have seen daylight on a land surface long ago, and sunk again to be covered by newer deposits, perhaps even a second or a third time, before they rose for the time that is the present. Yet all these profound changes took place so slowly that had we been living then we could have felt no motion, just as we feel no motion to-day, though the land is continuing to change all around us. The great alternations between land and water over large areas mark out to some extent the main periods tabulated on p. 34, for after each great submersion the rising land seems to have harboured plants and animals with somewhat different characters from those which inhabited it before. Similarly, when the next submersion laid down more rocks of limestone and sandstone, they enclosed the shells of some creatures different from those which had inhabited the seas of the region previously.

Through all the periods the actual rocks formed are very similar—shales, limestones, sandstones, clays. When any rocks happen to have preserved neither plant nor animal remains it is almost impossible to tell to which epoch they belong, except from a comparative study of their position as regards other rocks which do retain fossils. This depends on the fact that the physical processes of rock building have gone on throughout the history of the globe on very much the same lines as they are following at present. By the sifting power of water, fine mud, sand, pebbles, and other débris are separated from each other and collected in masses like to like. The fine mud will harden into shales, sandgrains massed together harden into sandstones, and so on, and when, after being raised once more to form dry land, they are broken up by wind and rain and brought down again to the sea, they settle out once again in a similar way and form new shales and sandstones; and so on indefinitely. But meantime the living things, both plant and animal, have been changing, growing, evolving, and the leafy twig brought down with the sandgrains in the flooded river of one epoch differs from that brought down by the river of a succeeding epoch—though it might chance that the sandgrains were the same identical ones. And hence it is by the remains of the plants and animals in a rock that we can tell to which epoch it belonged. Unless, of course, ready-formed fossils from an earlier epoch get mixed with it, coming as pebbles in the river in flood—but that is a subtle point of geological importance which we cannot consider here. Such cases are almost always recognizable, and do not affect the main proposition.

From the various epochs, the plants which have been preserved as fossils are in nearly all cases those which had lived on the land, or at least on swamps and marshes by the land. Of water plants in the wide sense, including both those growing in fresh water and those in the sea, we have comparatively few. This lack is particularly remarkable in the case of the seaweeds, because they were actually growing in the very medium in which the bulk of the rocks were formed, and which we know from recent experiments acts as a preservative for the tissues of land plants submerged in it. It must be remembered, however, that almost all the plants growing in water have very soft tissues, and are usually of small size and delicate structure as compared with land plants, and thus would stand less chance of being preserved, and would also stand less chance of being recognized to-day were they preserved. The mark on a stone of the impression of a soft film of a waterweed would be very slight as compared with that left by a leathery leaf or the woody twig of a land plant.

There are, of course, exceptions, and, as will be noted later on (see Chapter XVII), there are fossil seaweeds and fossil freshwater plants, but we may take it on the whole that the fossils we shall have to deal with and that give important evidence, are those of the land which had drifted out to sea, in the many cases when they are found in rocks together with sea shells.

Let us now consider very shortly the salient features of the seven epochs we have named as the chief divisions of time. The vegetation of the Carboniferous Period is better known to us than that of any other period except that of the present day, so that it will form the best starting-point for our consideration.

At this period there were, as there are to-day, oceans and continents, high lands, low lands, rivers and lakes, in fact, all the physical features of the present-day world, but they were all in different places from those of to-day. If we confine our attention to Britain, we find that at that period the far north, Scotland, Wales, and Charnwood were higher land, but the bulk of the southern area was covered by flat swamps or shallow inlets, where the land level gradually changed, slowly sinking in one place and slowly rising in others, which later began also to sink. Growing on this area wherever they could get a foothold were many plants, all different from any now living. Among them none bore flowers. A few families bore seeds in a peculiar way, differing widely from most seed-bearing plants of to-day. The most prevalent type of tree was that of which a stump is represented in the frontispiece, and of which there were many different species. These plants, though in size and some other ways similar to the great trees of to-day, were fundamentally different from them, and belonged to a very primitive family, of which but few and small representatives now exist, namely the Lycopods. Many other great trees were like hugely magnified “horsetails” or Equisetums; and there were also seed-bearing Gymnosperms of a type now extinct. There were ferns of many kinds, of which the principal ones belong to quite extinct families, as well as several other plants which have no parallel among living ones. Hence one may judge that the vegetation was rich and various, and that, as there were tall trees with seeds, the plants were already very highly evolved. Indeed, except for the highest group of all, the flowering plants, practically all the main groups now known were represented. The flora of the Devonian was very similar in essentials.

If that be so, it may seem unsatisfactory to place all the preceding æons under one heading, the Older Palæozoic. And, indeed, it is very unsatisfactory to be forced to do so. We know from the study of animal fossils that this time was vast, and that there were several well-defined periods in it during which many groups of animals evolved, and became extinct after reaching their highest development; but of the plants we know so little that we cannot make any divisions of time which would be of real value in helping us to understand them.

Fossil plants from the Early Palæozoic there are, but extremely few as compared with the succeeding period, and those few but little illuminative. In the later divisions of the Pre-Carboniferous some of the plants seem to belong to the same genera as those of the Carboniferous period. There is a fern which is characteristic of one of the earlier divisions, and there are several rather indefinite impressions which may be considered as seaweeds. There is evidence also that even one of the higher groups bearing seeds (the _Cordaiteæ_) was in full swing long before the Carboniferous period began. Hence, though of Older Palæozoic plants we know little of actual fact, we can surmise the salient truths; viz., that in that period those plants must have been evolving which were important in the Devonian and Carboniferous periods; that in the earlier part of that period they did not exist, and the simpler types only clothed the earth; and that further back still, even the simpler types had not yet evolved.

Names have been given to many fragmentary bits of fossils, but for practical purposes we might as well be without them. For the present the actual plants of the Older Palæozoic must remain in a misty obscurity, their forms we can imagine, but not know.

On the other hand, of the more recent periods, those succeeding the Carboniferous, we have a little more knowledge. Yet for all these periods, even the Tertiary immediately preceding the present day, our knowledge is far less exact and far less detailed than it is for that unique period, the Carboniferous itself.

The characteristic plants of the Carboniferous period are all very different from those of the present, and every plant of that date is now extinct. In the succeeding periods the main types of vegetation changed, and with each succeeding change advanced a step towards the stage now reached.

The Permian, geologically speaking, was a period of transition. Toward the close of the Carboniferous there were many important earth movements which raised the level of the land and tended to enclose the area of water in what is now Eastern Europe, and to make a continental area with inland seas. Many of the Carboniferous genera are found to extend through the Permian and then die out, while at the same time others became quite extinct as the physical conditions changed. The seed-bearing plants became relatively more important, and though the genus _Cordaites_ died out at the end of the period it was succeeded by an increasing number of others of more advanced type.

When we come to the older Mesozoic rocks, we have in England at any rate an area which was slowly submerging again. The more important of the plants which are preserved, and they are unfortunately all too few, are of a type which has not yet appeared in the earlier rocks, and are in some ways like the living _Cycas_, though they have many characters fundamentally different from any living type. In the vegetation of this time, plants of Cycad-like appearance seem to have largely predominated, and may certainly be taken as the characteristic feature of the period. The great Lycopod and Equisetum-like trees of the Carboniferous are represented now only by smaller individuals of the same groups, and practically all the genera which were flourishing in the Carboniferous times have become extinct.

The Cycad-like plants, however, were far more numerous and varied in character and widely spread than they ever were in any succeeding time. Still, no flowers (as we understand the word to-day) had appeared, or at least we have no indication in any fossil hitherto discovered, that true flowers were evolved until towards the end of the period (see, however, Chapter X).

The newer Mesozoic or Upper Cretaceous period represents a relatively deep sea area over England, and the rocks then formed are now known as the chalk, which was all deposited under an ocean of some size whose water must have been clear, and on the whole free from ordinary débris, for the chalk is a remarkably homogeneous deposit. From the point of view of plant history, the Upper Mesozoic is notable, because in it the flowering plants take a suddenly important position. Beds of this age (though of very different physical nature) are known all over the world, and in them impressions of leaves and fruits, or their casts, are well represented. The leaves are those of both Monocotyledons and Dicotyledons, and the genera are usually directly comparable with those now living, and sometimes so similar that they appear to belong to the same genus. The cone-bearing groups of the Gymnosperms are still present and are represented by a number of forms, but they are far fewer in varieties than are the groups of flowering plants—while the Cycad-like plants, so important in the Lower Mesozoic, have relatively few representatives. There is, it almost seems, a sudden jump from the flowerless type of vegetation of the Lower Mesozoic, to a flora in the Upper Mesozoic which is strikingly like that of the present day.

The Tertiary period is a short one (geologically speaking, and compared with those going before it), and during it the land level rose again gradually, suffering many great series of earth movements which built most of the mountain chains in Europe which are standing to the present day. In the many plant-containing deposits of this age, we find specimens indicating that the flora was very similar to the plants now living, and that flowering plants held the dominant position in the forests, as they do to-day. In fact, from the point of view of plant evolution, it is almost an arbitrary and unnecessary distinction to separate the Tertiary epoch from the present, because the main features of the vegetation are so similar. There are, however, such important differences in the distribution of the plants of the Tertiary and those of the present times, that the distinction is advisable; but it must always be remembered that it is not comparable with the wide differences between the other epochs.

Among the plants now living we find representatives of most, though not of all, of the great _groups_ of plants which have flourished in the past, though in the course of time all the species have altered and those of the earliest earth periods have become extinct. The relative importance of the different groups changes greatly in the various periods, and as we proceed through the ages of time we see the dominant place in the plant world held successively by increasingly advanced types, while the plants which dominated earlier epochs dwindle and take a subordinate position. For example, the great trees of the Carboniferous period belonged to the Lycopod family, which to-day are represented by small herbs creeping along the ground. The Cycad-like plants of the Mesozoic, which grew in such luxuriance and in such variety, are now restricted to a small number of types scattered over the world in isolated localities.

During all the periods of which we have any knowledge there existed a rich and luxuriant vegetation composed of trees, large ferns, and small herbs of various kinds, but the members of this vegetation have changed fundamentally with the changing earth, and unlike the earth in her rock-forming they have never repeated themselves.

Chapter V Stages In Plant Evolution

To attempt any discussion of the _causes_ of evolution is far beyond the scope of the present work. At present we must accept life as we find it, endowed with an endless capacity for change and a continuous impulse to advance. We can but study in some degree the _course_ taken by its changes.

From the most primitive beginnings of the earliest periods, enormous advance had been made before we have any detailed records of the forms. Yet there remain in the world of to-day numerous places where the types with the simplest structure can still flourish, and successfully compete with higher forms. Many places which, from the point of view of the higher plants, are undesirable, are well suited to the lower. Such places, for example, as the sea, and on land the small nooks and crannies where water drops collect, which are useless for the higher plants, suffice for the minute forms. In some cases the lower plants may grow in such masses together as to capture a district and keep the higher plants from it. Equisetum (the horsetail) does this by means of an extensive system of underground rhizomes which give the plant a very strong hold on a piece of land which favours it, so that the flowering plants may be quite kept from growing there.

In such places, by a variety of means, plants are now flourishing on the earth which represent practically all the main stages of development of plant life as a whole. It is to the study of the simpler of the living forms that we owe most of our conceptions of the course taken by evolution. Had we to depend on fossil evidence alone, we should be in almost complete ignorance of the earliest types of vegetation and all the simpler cohorts of plants, because their minute size and very delicate structure have always rendered them unsuitable for preservation in stone. At the same time, had we none of the knowledge of the numerous fossil forms which we now possess, there would be great gaps in the series which no study of living forms could supply. It is only by a study and comparison of both living and fossil plants of all kinds and from beds of all ages that we can get any true conception of the whole scheme of plant life.

Grouping together all the main families of plants at present known to us to exist or to have existed, we get the following series:—

Group. Common examples of typical families in the group.

Thallophyta Algæ Seaweeds. Fungi Moulds and toadstools. Bryophyta Hepaticæ Liverworts. Musci Mosses. Pteridophyta Equisetales Horsetails. Sphenophyllales* fossil only, _Sphenophyllum_. Lycopodales Club-moss. Filicales Bracken fern. Pteridospermæ Lyginodendræ* fossil only, _Sphenopteris_. Gymnosperms Cycadales Cycads. Bennettitales* fossil only, _Bennettites_. Ginkgoales Maidenhair Tree. Cordaitales* fossil only, _Cordaites_. Coniferales Pine, Yew. Gnetales Welwitschia. Angiosperms Monocotyledons Lily, Palm, Grass. Dicotyledons Rose, Oak, Daisy.

In this table the different groups have not a strictly equivalent scientific value, but each of those in the second column represents a large and well-defined series of primary importance, whose members could not possibly be included along with any of the other groups.

Those marked with an asterisk are known only as fossils, and it will be seen that of the seventeen groups, so many as four are known only in the fossil state. This indicates, however, but a part of their importance, for in nearly every other group are many families or genera which are only known as fossils, though there are living representatives of the group as a whole.

In this table the individual families are not mentioned, because for the present we need only the main outline of classification to illustrate the principal facts about the course of evolution. As the table is given, the simplest families come first, the succeeding ones gradually increasing in complexity till the last group represents the most advanced type with which we are acquainted, and the one which is the dominant group of the present day.

This must not be taken as a suggestion that the members of this series have evolved directly one from the other in the order in which they stand in the table. That is indeed far from the case, and the relations between the groups are highly complex.

It must be remarked here that it is often difficult, even impossible, to decide which are the most highly evolved members of any group of plants. Each individual of the higher families is a very complicated organism consisting of many parts, each of which has evolved more or less independently of the others in response to some special quality of the surroundings. For instance, one plant may require, and therefore evolve, a very complex and well-developed water-carriage system while retaining a simple type of flower; another may grow where the water problem does not trouble it, but where it needs to develop special methods for getting its ovules pollinated; and so on, in infinite variety. As a result of this, in almost all plants we have some organs highly evolved and specialized, and others still in a primitive or relatively primitive condition. It is only possible to determine the relative positions of plants on the scale of development by making an average conclusion from the study of the details of all their parts. This, however, is beset with difficulties, and in most cases the scientist, weighed by personal inclinations, arbitrarily decides on one or other character to which he pays much attention as a criterion, while another scientist tends to lay stress on different characters which may point in another direction.

In no group is this better illustrated than among the Coniferæ, where the relative arrangement of the different families included in it is still very uncertain, and where the observations of different workers, each dealing mainly with different characters in the plants, tend to contradict each other.

This, however, as a byword. Notwithstanding these difficulties, which it would be unfair to ignore, the main scheme of evolution stands out clearly before the scientist of to-day, and his views are largely supported by many important facts from both fossil and living plants.

Very strong evidence points to the conclusion that the most primitive plants of early time were, like the simplest plants of to-day, water dwellers. Whether in fresh water or the sea is an undecided point, though opinion seems to incline in general to the view that the sea was the first home of plant life. It can, however, be equally well, and perhaps even more successfully argued, that the freshwater lakes and streams were the homes of the first families from which the higher plants have gradually been evolved.

For this there is no direct evidence in the rocks, for the minute forms of the single soft cells assumed by the most primitive types were just such as one could not expect to be successfully fossilized. Hence the earliest stages must be deduced from a comparative study of the simplest plants now living. Fortunately there is much material for this in the numerous waters of the earth, where swarms of minute types in many stages of complexity are to be found.

The simplest type of plants now living, which appears to be capable of evolution on lines which might have led to the higher plants, is that found in various members of the group of the Protococcoideæ among the Algæ. The claim of bacteria and other primitive organisms of various kinds to the absolute priority of existence is one which is entirely beyond the scope of a book dealing with fossil plants. The early evolution of the simple types of the Protococcoideæ is also somewhat beyond its scope, but as they appear to lie on the most direct “line of descent” of the majority of the higher plants it cannot be entirely ignored. From the simpler groups of the green Algæ other types have specialized and advanced along various directions, but among them there seems an inherent limitation, and none but the protococcoid forms seem to indicate the possibility of really high development.

[Illustration: Fig. 17.—A Protococcoid Plant consisting of one cell

_p_, Protoplasm; _n_, nucleus; _g_, colouring body or chloroplast; _w_, cell wall.]

In a few words, a typical example of one of the simple Protococcoideæ may be described as consisting of a mass of protoplasm in which lie a recognizable nucleus and a green colouring body or chloroplast, with a cell wall or skin surrounding these vital structures, a cell wall that may at times be dispensed with or unusually thickened according as the need arises. This plant is represented in fig. 17 in a somewhat diagrammatic form.

In such a case the whole plant consists of one single cell, living surrounded by the water, which supplies it with the necessary food materials, and also protects it from drying up and from immediate contact with any hard or injurious object. When these plants propagate they divide into four parts, each one similar to the original cell, which all remain together within the main cell wall for a short time before they separate.

If now we imagine that the four cells do not separate, but remain together permanently, we can see the possibility of a beginning of specialization in the different parts of the cell. The single living cell is equally acted on from all sides, and in itself it must perform all the life functions; but where four lie together, each of the four cells is no longer equally acted on from all sides. This shows clearly in the diagram of a divided cell given in fig. 18. Here it is obvious that one side of each of the four cells, viz. that named _a_ in the diagram, is on the outside and in direct contact with the water and external things; but walls _b_ and _c_ touch only the corresponding walls of the neighbouring cells. Through walls _b_ and _c_ no food and water can enter directly, but at the same time they are protected from injury and external stimulus. Hence, in this group of four cells there is a slight differentiation of the sides of the cells. If now we imagine that each of the four cells, still remaining in contact, divides once more into four members, each of which reaches mature size while all remain together, then we have a group of sixteen cells, some of which will be entirely inside, and some of which will have walls exposed to the environment.

[Illustration: Fig. 18.—Diagram of Protococcoid Cell divided into four daughter cells. Walls _a_ are external, and walls _b_ and _c_ in contact with each other.]

If the cells of the group all divide again, in the manner shown the mass will become more than one cell thick, and the inner cells will be more completely differentiated, for they will be entirely cut off from the outside and all direct contact with water and food materials, and will depend on what the outer cells transmit to them. The outer cells will become specialized for protection and also for the absorption of the water and salts and air for the whole mass. From such a plastic group of green cells it is probable that the higher and increasingly complex forms of plants have evolved. There are still living plants which correspond with the groups of four, sixteen, &c., cells just now theoretically stipulated.

[Illustration: Fig. 19.—A, Details of Part of the Tissues in a Stem of a Flowering Plant. B, Diagram of the Whole Arrangement of Cross Section of a Stem: _e_, Outer protecting skin; _g_, green cells; _s_, thick-walled strengthening cells; _p_, general ground tissue cells. V, Groups of special conducting tissues: _x_, vessels for water carriage; _px_, first formed of the water vessels; _c_, growing cells to add to the tissues; _b_, food-conducting cells; _ss_, strengthening cells.]

The higher plants of to-day all consist of very large numbers of cells forming tissues of different kinds, each of which is specialized more or less, some very elaborately, for the performance of certain functions of importance for the plant body as a whole. With the increase in the number of cells forming the solid plant body, the number of those living wholly cut off from the outside becomes increasingly great in comparison with those forming the external layer. Some idea of the complexity and differentiation of this cell mass is given in fig. 19, A, which shows the relative sizes and shapes of the cells composing a small part of the stem of a common flowering plant. The complete section would be circular and the groups V would be repeated round it symmetrically, and the whole would be enclosed by an unbroken layer of the cells marked _e_, as in the diagram B.

[Illustration: Fig. 20.—Conducting Cells and Surrounding Tissue seen in fig. 19, A, cut lengthways. _px_, First formed vessels for water conduction; _x_, larger vessel; _b_, food-conducting cells; _ss_, strengthening cells; _p_, general ground tissue.]

In the tissues of the higher plants the most important feature is the complex system of conducting tissues, shown in the young condition in V in fig. 19, A. In them the food and water conducting elements are very much elongated and highly specialized cells, which run between the others much like a system of pipes in the brickwork of a house. These cells are shown cut longitudinally in fig. 20, where they are lettered to correspond with the cells in fig. 19, A, with which they should be compared. In such a view the great difference between the highly specialized cells _x_, _px_, _b_, &c., and those of the main mass of ground tissue _p_ becomes apparent.

Even in the comparatively simply organized groups of the Equisetales and Lycopodiales the differentiation of tissues is complete. In the mosses, and still more in the liverworts, it is rudimentary; but they grow in very damp situations, where the conduction of water and the protection from too much drying is not a difficult problem for them. As plants grow higher into the air, or inhabit drier situations, the need of specialization of tissues becomes increasingly great, for they are increasingly liable to be dried, and therefore need a better flow of water and a more perfect protective coat.

It is needless to point out how the individual cells of a plant, such as that figured in figs. 19 and 20, have specialized away from the simple type of the protococcoid cell in their mature form. In the young growing parts of a plant, however, they are essentially like protococcoid cells of squarish outline, fitting closely to each other to make a solid mass, from which the individual types will differentiate later and take on the form suitable for the special part they have to play in the economy of the whole plant.

To trace the specialization not only of the tissues but of the various parts of the whole plant which have become elaborate organs, such as leaves, stems, and flowers, is a task quite beyond the present work to attempt. From the illustrations given of tissue structure from plants at the two ends of the series much can be imagined of the inevitable intermediate stages in tissue evolution.

As regards the elaboration of organs, and particularly of the reproductive organs, details will be found throughout the book. In judging of the place of any plant in the scale of evolution it is to the reproductive organs that we look for the principal criteria, for the reproductive organs tend to be influenced less by their physical surroundings than the vegetative organs, and are therefore truer guides to natural relationships.

In the essential cells of the reproductive organs, viz. the egg cell and the male cell, we get the most primitively organized cells in the plant body. In the simpler families both male and female cells return to the condition of a free-swimming protococcoid cell, and in all but the highest families the male cell requires a liquid environment, in which it _swims_ to the egg cell. In the higher families the necessary water is provided within the structure of the seed, and the male cell does not swim, a naked, solitary cell, out into the wide world, as it does in all the families up to and including the Filicales. In the Coniferæ and Angiosperms the male cell does not swim, but is passive (or largely so), and is brought to the egg cell. One might almost say that the whole evolution of the complex structures found in fruiting cones and flowers is a result of the need of protection of the delicate, simple reproductive cells and the embryonic tissues resulting from their fusion. The lower plants scatter these delicate cells broadcast in enormous numbers, the higher plants protect each single egg cell by an elaborate series of tissues, and actually bring the male cell to it without ever allowing either of them to be exposed.

It must be assumed that the reader possesses a general acquaintance with the living families tabulated on p. 44; those of the fossil groups will be given in some detail in succeeding chapters which deal with the histories of the various families. It is premature to attempt any general discussion of the evolution of the various groups till all have been studied, so that this will be reserved for the concluding chapters.

Chapter VI Minute Structure Of Fossil Plants—Likenesses To Living Ones

The individual plants of the Coal Measure period differed entirely from those now living; they were more than merely distinct species, for in the main even the families were largely different from the present ones. Nevertheless, when we come to examine the minute anatomy of the fossils, and the cells of which they are composed, we find that between the living and the fossil cell types the closest similarity exists.

From the earliest times of which we have any knowledge the elements of the plant body have been the same, though the types of structures which they built have varied in plan. Individual _cells_ of nearly every type from the Coal Measure period can be identically matched with those of to-day. In the way the walls thickened, in the shapes of the wood, strengthening or epidermal cells, in the form of the various tissues adapted to specific purposes, there is a unity of organization which it is reasonable to suppose depends on the fundamental qualities inherent in plant life.

This will be illustrated best, perhaps, by tabulating the chief modifications of cells which are found in plant tissues. The illustrations of these types in the following table are taken from living plants, because from them figures of more diagrammatic clearness can be made, and the salient characters of the cells more easily recognized. Comparison of these typical cells with those illustrated from the fossil plants reveals their identity in essential structure, and most of them will be found in the photos of fossils in these pages, though they are better recognized in the actual fossils themselves.

Principal Types of Plant Cells

Epidermal.

[Illustration: Fig. 21

_Epidermis._—Protecting layer or skin. Cells with outer wall thickened in many cases (fig. 21, _a_ and _b_). Compare fossil epidermis in fig. 34, _e_.]

[Illustration: Fig. 22 _Hairs._—Extensions of epidermis cells. Single cells, or complex, as fig. 22, _h_, where _e_ is epidermis and _p_ parenchyma. Compare fossil hairs in figs. 79 and 120.]

[Illustration: Fig. 23

_Stomates._—Breathing pores in the epidermis. Seen in surface view as two-lipped structures (fig. 23). _s_, Stomates; _e_, epidermis cells. Compare fossil stomates in fig. 8.]

Ground Tissue

[Illustration: Fig. 24

_Parenchyma._—Simple soft cells, either closely packed, as in fig. 24, or with air spaces between them. Compare 78, B, for fossil.]

[Illustration: Fig. 25

_Palisade._—Elongated, closely packed cells, _p_, chiefly in leaves, lying below the epidermis, _e_, fig. 25. Compare fig. 34, _p_, for fossil palisade.]

[Illustration: Fig. 26

_Endodermis._—Cells with specially thickened walls, _en_, lying as sheath between the parenchyma, _c_, of ground tissue, and the vascular tissue, _s_, fig. 26. Compare fig. 108 for fossil endodermis.]

[Illustration: Fig. 27

_Latex cells._—Large, often much elongated cells, _m_, lying in the parenchyma, _p_, fig. 27, which are packed with contents. Compare fig. 107, _s_.]

[Illustration: Fig. 28

_Sclerenchyma._—Thick-walled cells among parenchyma for strengthening, fig. 28. Compare fig. 34, _s_.]

[Illustration: Fig. 29

_Cork._—Layers of cells replacing the epidermis in old stems. Outer cells, _o_, crushed; _k_, closely packed cork cells; stone cells, _s_, fig. 29. Compare fig. 95, _k_.

_Cork cambium._—Narrow, actively dividing cells, _c_ in fig. 29, giving rise to new cork cells in consecutive rows.]

[Illustration: Fig. 30

_Tracheides._—Specially thickened cells in the parenchyma, usually for water storage, _t_, fig. 30. Compare fig. 95, _t_.]

Vascular Tissue

[Illustration: Fig. 31

_Wood._—_Protoxylem_, tracheids and vessels, long, narrow elements, with spiral or ring-like thickenings, _s_^1 and _s_^2, fig. 31. Compare fig. 81, A, _px_, for fossil.

_Metaxylem_, long elements, tracheids and vessels. Some with narrow pits, as _t_ in fig. 31; others with various kinds of pits. In transverse section seen in fig. 33, w, fossil in fig. 114, _w_.

_Wood parenchyma._—Soft cells associated with the wood, _p_ in fig. 31. Fossil in fig. 81, B, _p_.

_Wood sclerenchyma._—Hard thickened cells in the wood.]

[Illustration: Fig. 32

_Bast._—_Sieve tubes_, long cells which carry foodstuffs, cross walls pitted like sieves, _s_, fig. 32. In transverse section in fig. 33.

_Companion cells_, narrow cells with rich proteid contents, _c_, fig. 32. In transverse section at _c_, fig. 33.

_Bast parenchyma._—Soft unspecialized cells mixed with the sieve tubes, _p_, fig. 32.

_Bast fibres._—Thick-walled sclerenchymatous cells mixed with, or outside, the soft bast.]

[Illustration: Fig. 33

_Cambium._—Narrow cells, like those of the cork cambium, which lie between the wood and bast, and give rise to new tissues of each kind, _cb_, fig. 33. Compare fig. 114, fossil.]

There are, of course, many minor varieties of cells, but these illustrate all the main types.

Among the early fossils, however, one type of wood cell and one type of bast cell, so far as we know, are not present. These cells are the true _vessels_ of the wood of flowering plants, and the long bast cells with their companion proteid cells. The figure of a metaxylem wood cell, shown in fig. 31, _t_, shows the more primitive type of wood cell, which has an oblique cross wall. This type of wood cell is found in all the fossil trees, and all the living plants except the flowering plants. The vessel type, which is that in the big wood vessels of the flowering plants, and has no cross wall, is seen in fig. 20, _x_.

The similarity between the living cells and those of the Coal Measure fossils is sufficiently illustrated to need no further comment. This similarity is an extremely helpful point when we come to an interpretation of the fossils. In living plants we can study the physiology of the various kinds of cells, and can deduce from experiment exactly the part they play in the economy of the whole plant. From a study of the tissues in any plant structure we know what function it performed, and can very often estimate the nature of the surrounding conditions under which the plant was growing. To take a single example, the palisade tissue, illustrated in fig. 25, _p_, in living plants always contains green colouring matter, and lies just below the epidermis, usually of leaves, but sometimes also of green stems. These cells do most of the starch manufacture for the plant, and are found best developed when exposed to a good light. In very shady places the leaves seldom have this type of cell. Now, when cells just like these are found in fossils (as is illustrated in fig. 34), we can assume all the physiological facts mentioned above, and rest assured that that leaf was growing under normal conditions of light and was actively engaged in starch-building when it was alive. From the physiological standpoint the fossil leaf is entirely the same as a normal living one.

[Illustration: Fig. 34.—From a Photo of a Fossil Lea

_e_, Epidermis; _p_, palisade cells; _pr_, soft parenchyma cells (poorly preserved); _s_, sclerenchyma above the vascular bundle.]

From the morphological standpoint, also, the features of the plant body from the Coal Measure period fall into the same divisions as those of the present. Roots, stems, leaves, and reproductive organs, the essentially distinct parts of a plant, are to be found in a form entirely recognizable, or sufficiently like that now in vogue to be interpreted without great difficulty. In the detailed structure of the reproductive organs more changes have taken place than in any others, both in internal organization and external appearance.

Already, in the Early Palæozoic period, the distinction between leaves, stems, roots, and reproductive organs was as clearly marked as it is to-day, and, judging by their structure, they must each have performed the physiological functions they now do. Roots have changed least in the course of time, probably because, in the earth, they live under comparatively uniform conditions in whatever period of the world’s history they are growing. Naturally, between the roots of different species there are slight differences; but the likeness between fern roots from the Palæozoic and from a living fern is absolutely complete. This is illustrated in fig. 35, which shows the microscopic structure of the two roots when cut in transverse direction. The various tissues will be recognized as coming into the table on p. 54, so that both in the details of individual cells and in the general arrangement of the cell groups or tissues the roots of these fossil and living ferns agree.

[Illustration: Fig. 35.—A, Root of Living Fern. B, Root of Palæozoic Fossil Fern. _c_, Cortex; _px_, protoxylem in two groups; _m_, metaxylem; _s_, space in fossil due to decay of soft cells.]

Among stems there has been at all periods more variety than among the roots of the corresponding plants, and in the following chapter, when the differences between living and fossil plants will be considered, there will be several important structures to notice. Nevertheless, there are very many characters in which the stems from such widely different epochs agree. The plants in the palæozoic forests were of many kinds, and among them were those with weak trailing stems which climbed over and supported themselves on other plants, and also tall, sturdy shafts of woody trees, many of which were covered with a corky bark. Leaves were attached to the stems, either directly, as in the case of some living plants, or by leaf stalks. In external appearance and in general function the stems then were as stems are now. In the details of the individual cells also the likeness is complete; it is in the grouping of the cells, the anatomy of the tissues, that the important differences lie. It has been remarked already that increase in complexity of the plant form usually goes with an increase in complexity of the cells and variety of the tissues. The general ground tissue in nearly all plants is very similar; it is principally in the vascular system that the advance and variety lie.

Plant anatomists lay particular stress on the vascular system, which, in comparison with animal anatomy, holds an even more important position than does the skeleton. To understand the essential characters of stems, both living and fossil, and to appreciate their points of likeness or difference, it is necessary to have some knowledge of the general facts of anatomy; hence the main points on which stress is laid will be given now in brief outline.

Leaving aside consideration of the more rudimentary and less defined structure of the algæ and mosses, all plants may be said to possess a “vascular system”. This is typically composed of elongated wood (or xylem) with accessory cells (see p. 57, table), and bast (phloem), also with accessory cells. These specialized conducting elements lie in the ground tissue, and in nearly all cases are cut off from direct contact with it by a definite sheath, called the endodermis (see p. 55, fig. 26). Very often there are also groups or rings of hard thick-walled cells associated with the vascular tissues, which protect them and play an important part in the consolidation of the whole stem.

[Illustration: Fig. 36.—Diagram of Simplest Arrangement of Complete Stele in a Stem

W, Central solid wood; P, ring of bast; E, enclosing sheath of endodermis; C, ground tissue or cortex.]

The simplest, and probably evolutionally the most primitive form which is taken by the vascular tissues, is that of a single central strand, with the wood in the middle, the bast round it, and a circular endodermis enclosing all, as in fig. 36, which shows a diagram of this arrangement. Such a mass of wood and bast surrounded by an endodermis, is technically known as a _stele_, a very convenient term which is much used by anatomists. In its simplest form (as in fig. 36) it is called a _protostele_, and is to be found in both living and fossil plants. A number of plants which get more complex steles later on, have protosteles in the early stages of their development, as in _Pteris aurita_ for example, a species allied to the bracken fern, which has a hollow ring stele when mature.

[Illustration: Fig. 37.—Diagram of a Stele with a few Cells of Pith _p_ in the Middle of the Wood. Lettering as in fig. 36]

[Illustration: Fig. 38.—Diagram showing Extensive Pith _p_ in the Wood. Lettering as in fig. 36]

The next type of stele is quite similar to the protostele, but with the addition of a few large unspecialized cells in the middle of the wood (_p_, fig. 37); these are the commencement of the hollowing process which goes on in the wood, resulting later in the formation of a considerable pith, as is seen in fig. 38, where the wood is now a hollow cylinder, as the phloem has been from the first. When this is the case, a second sheath or endodermis generally develops on the inner side of the wood, outside the pith, and cuts the vascular tissues off from the inner parenchyma. A further step is the development of an inner cylinder of bast so that the vascular ring is completely double, with endodermis on both sides of the cylinder, as is seen in fig. 39.

[Illustration: Fig. 39.—A Cylindrical Stele, with _e_, inner endodermis, and _ph_, inner phloem; W, wood; P, outer phloem; E, outer endodermis. L, part of the stele going out to supply a large leaf, thus breaking what would otherwise appear as a closed ring stele]

In all these cases there is but one strand or cylinder, of vascular tissue in the stem, but one stele, and this type of anatomy is known as the _monostelic_ or single-steled type.

[Illustration: Fig. 40.—A Ring Stele apparently broken up into a Number of Protosteles by many Leaf Gaps]

When from the double cylinder just described a strand of tissue goes off to supply a large leaf, a considerable part of the stele goes out and breaks the ring. This is shown in fig. 39, where L is the part of the stele going to a leaf, and the rest the broken central cylinder. When the stem is short, and leaves grow thickly so that bundles are constantly going out from the main cylinder, this gets permanently broken, and its appearance when cut across at any given point is that of a group of several steles arranged in a ring, each separate stele being like the simple protostele in its structure. See fig. 40. This type of stem has long been known as _polystelic_ (_i.e._ many-steled), and it is still a convenient term to describe it by. There has been much theoretical discussion about the true meaning of such a “polystelic” stem, which cannot be entered into here; it may be noted, however, that the various strands of the broken ring join up and form a meshwork when we consider the stem as a whole, it is only in a single section that they appear as quite independent protosteles. Nevertheless, as we generally consider the anatomy of stems in terms of single sections, and as the descriptive word “polystelic” is a very convenient and widely understood term, it will be used throughout the book when speaking of this type of stem anatomy.

Such a type as this, shown in fig. 40, is already complex, but it often happens that the steles branch and divide still further, until there is a highly complicated and sometimes bewildering system of vascular strands running through the ground tissue in many directions, but cut off from it by their protective endodermal sheaths. Such complex systems are to be found both in living and fossil plants, more especially in many of the larger ferns (see fig. 88).

Higher plants in general, however, and in particular flowering plants, do not have a polystelic vascular arrangement, but a specialized type of monostele.

[Illustration: Fig. 41.—Monostele in which the Central Pith is Star-shaped, and the Wood breaking up into Separate Groups

_p_, Pith; W, wood; P, phloem; E, endodermis; C, cortex.]

Referring again to fig. 37 as a starting-point, imagine the pith in the centre to spread in a star-shaped form till the points of the star touched the edges of the ring, and thus to break the wood ring into groups. A stage in this process (which is not yet completed) is shown in fig. 41, while in fig. 42 the wood and bast groups are entirely distinct. In the flowering plants the cells of the endodermis are frequently poorly characterized, and the pith cells resemble those of the cortical ground tissue, so that the separate groups of wood and bast (usually known as “vascular bundles”, in distinction from the “steles” of fig. 40) appear to lie independently in the ground tissue. These strands, however, must not be confused with steles, they are only fragments of the single apparently broken up stele which runs in the stem.

[Illustration: Fig. 42.—Monostele in which the Pith has invaded all the Tissues as far as the Endodermis, and broken the Wood and Phloem up into Separate Bundles. These are usually called “vascular bundles” in the flowering plants]

[Illustration: Fig. 43.—Showing actively growing Zone _c_ (Cambium) in the Vascular Bundles, and joining across the ground tissue between them]

The vascular bundle, of all except the Monocotyledons, has a potentiality for continued growth and expansion which places it far above the stele in value for a plant of long life and considerable growth. The cells lying between the wood and the bast, the soft parenchyma cells always accompanying such tissues, retain their vitality and continue to divide with great regularity, and to give rise to a continuous succession of new cells of wood on the one side and bast on the other; see fig. 33, _c_, _b_. In this way the primary, distinct vascular bundles are joined by a ring of wood, see fig. 43, to which are added further rings every season, till the mass of wood becomes a strong solid shaft. This ever-recurring activity of the cambium gives rise to what are known as “annual rings” in stems, see fig. 44, in which the wood shows both primary distinct groups in the centre, and the rings of growth of later years.

Cambium with this power of long-continued activity is found in nearly all the higher plants of to-day (except the Monocotyledons), but in the fern and lycopod groups it is in abeyance. Certain cases from nearly every family of the Pteridophytes are known, where some slight development of cambium with its secondary thickening takes place, but in the groups below the Gymnosperms cambium has almost no part to play. On the other hand, so far back as the Carboniferous period, the masses of wood in the Pteridophyte trees were formed by cambium in just the same way as they are now in the higher forms. Its presence was almost universal at that time in the lower groups where to-day there are hardly any traces of it to be found.

[Illustration: Fig. 44.—Stem with Solid Cylinder of Wood developed from the Cambium, showing three “annual rings”. In the centre may still be seen the separate groups of the wood of the primary “vascular bundles”]

It will be seen from this short outline of the vascular system of plants, that there is much variety possible from modifications of the fundamental protostele. It is also to be noted that the plants of the Coal Measures had already evolved all the main varieties of steles which are known to us even now,[6] and that the development of secondary thickening was very widespread. In several cases the complexity of type exceeds that of modern plants (see Chap. VII), and there are to be found vascular arrangements no longer extant.

When we turn to the _Reproductive Organs_, we find that the points of likeness between the living and the fossil forms are not so numerous or so direct as they are in the case of the vegetative system.

[Illustration: Fig. 45.—Fern Sporangia

A, fossil; B, living.]

As has been indicated, the families of plants typical of the Coal Measures were not those which are the most prominent to-day, but belonged to the lower series of Pteridophytes. In their simpler forms the fructifications then and now resemble each other very closely, but in the more elaborate developments the points of variety are more striking, so that they will be dealt with in the following chapter. Cases of likeness are seen in the sporangia of ferns, some of which appear to have been practically identical with those now living. This is illustrated in fig. 45, which shows the outline of the cells of the sporangia of living and fossil side by side.

[Illustration: Fig. 46.—A, Living Lycopod cone; B, _Lepidodendron_ (fossil) cone. _a_, Axis; _s_, scale; S, sporangium with spores. One side of a longitudinal section]

In the general structure also of the cones of the simpler types of _Lepidodendron_ (fossil, see frontispiece) there is a close agreement with the living Lycopods, though as regards size and output of spores there was a considerable difference in favour of the fossils. The plan of each is that round the axis of the cone simple scales are arranged, on each of which, on its upper side, is seated a large sporangium bearing numerous spores all of one kind (see fig. 46).

Equally similar are the cones of the living Equisetum and some of the simple members of the fossil family Calamiteæ, but the more interesting cases are those where differences of an important morphological nature are to be seen.

As regards the second[7] generation there is some very important evidence, from extremely young stages, which has recently been given to the world. In a fern sporangium _germinating spores_ were fossilized so as to show the first divisions of the spore cell. These seem to be identical with the first divisions of some recent ferns (see fig. 47). This is not only of interest as showing the close similarity in detail between plants of such widely different ages, but is a remarkable case of delicate preservation of soft and most perishable structures in the “coal balls”.

[Illustration: Fig. 47.—Germinating Fern Spores

A and B, from carboniferous fossils; C, living fern. (A and B after Scott.)]

While these few cases illustrate points of likeness between the fructifications of the Coal Measures and of to-day, the large size and successful character of the primitive Coal Measure plants was accompanied by many developments on the part of their reproductive organs which are no longer seen in living forms, and the greater number of palæozoic fructifications must be considered in the next chapter.

Chapter VII Minute Structure Of Fossil Plants—Differences From Living Ones

We have seen in the last chapter that the main morphological divisions, roots, stems, leaves, and fructifications, were as distinct in the Coal Measure period as they are now. There is one structure, however, found in the Coal Measure fossils, which is hardly paralleled by anything similar in the living plants, and that is the fossil known as _Stigmaria_. _Stigmaria_ is the name given, not to a distinct species of plant, but to the large rootlike organs which we know to have belonged to all the species of _Lepidodendron_ and of _Sigillaria_. In the frontispiece these organs are well seen, and branch away at the foot of the trunk, spreading horizontally, to all appearance merely large roots. They are especially regularly developed, however, the main trunk giving rise always to four primary branches, these each dividing into two equal branches, and so on—in this they are unlike the usual roots of trees. They bore numerous rootlets, of which we know the structure very well, as they are the commonest of all fossils, but in their internal anatomy the main “roots” had not the structure which is characteristic of roots, but were like _stems_. In living plants there are many examples of stems which run underground, but they always have at least the rudiments of leaves in the form of scales, while the fossil structures have apparently no trace of even the smallest scales, but bear only rootlets, thus resembling true roots. The questions of morphology these structures raise are too complex to be discussed here, and Stigmaria is only introduced as an example, one of the very few available, of a palæozoic structure which seems to be of a nature not clearly determinable as either root, stem, leaf, or fructification. Among living plants the fine rootlike rhizophores of Selaginella bear some resemblance to Stigmaria in essentials, though so widely different from them in many ways, and they are probably the closest analogy to be found among the plants of to-day.

The individual cells, we have already seen, are strikingly similar in the case of fossil and living plants. There are, of course, specific varieties peculiar to the fossils, of which perhaps the most striking seem to be some forms of _hair_ cells. For example, in a species of fern from the French rocks there were multicellular hairs which looked like little stems of Equisetum owing to regular bands of teeth at the junctions of the cells. These hairs were quite characteristic of the species—but hairs of all sorts have always abounded in variety, so that such distinction has but minor significance.

[Illustration: Fig. 48.—Stele of _Lepidodendron_ W, surrounded by a small ring of secondary wood S]

As was noted in the table (p. 58) the only cell types of prime importance which were not evolved by the Palæozoic plants were the wood vessels, phloem and accompanying cells which are characteristic of the flowering plants.

Among the fossils the vascular arrangements are most interesting, and, as well as all the types of stele development noted in the previous chapter as common to both living and fossil plants, there are further varieties found only among the fossils (see fig. 50).

The simple protostele described (on p. 61) is still found, particularly in the very young stages of living ferns, but it is a type of vascular arrangement which is not common in the mature plants of the present day. In the Coal Measure period, however, the protostele was characteristic of one of the two main groups of ferns. In different species of these ferns, the protostele assumed a large variety of shapes and forms as well as the simple cylindrical type. The central mass of wood became five-rayed in some, star-shaped, and even very deeply lobed, with slightly irregular arms, but in all these cases it remained fundamentally monostelic. Frequently secondary tissue developed round the protosteles of plants whose living relatives have no such tissue. A case of this kind is illustrated in fig. 48, which shows a simple circular stele surrounded by a zone of secondary woody tissue in a species of _Lepidodendron_.

[Illustration: Fig. 49.—_Lepidodendron_, showing Part of the Hollow Ring of Primary Wood W, with a relatively large amount of Secondary Tissue S, surrounding it]

In many species of _Lepidodendron_ the quantity of secondary wood formed round the primary stele was very great, so that (as is the case in higher plants) the primary wood became relatively insignificant compared with it. In most species of _Lepidodendron_ the primary stele is a hollow ring of wood (cf. fig. 38, p. 62) round which the secondary wood developed, as is seen in fig. 49. These two cases illustrate a peculiarity of fossil plants. Among living ones the solid and the simple ring stele are almost confined to the Pteridophytes, where secondary wood does not develop, but the palæozoic Pteridophytes, while having the simple primary types of steles, had quantities of secondary tissue, which was correlated with their large size and dominant position.

[Illustration: Fig. 50.—Diagram of Steles of the English _Medullosa_, showing three irregular, solid, steles A, with secondary thickenings S, all round each. _a_, Small accessory steles]

Among _polystelic_ types (see p. 63) we find interesting examples in the fossil group of the _Medulloseæ_, which are much more complex than any known at present, both owing to their primary structure and also to the peculiar fact that all the steles developed secondary tissue towards the inner as well as the outer side. One of the simpler members of this family found in the English Coal Measures is illustrated in fig. 50. Here there are three principal protosteles (and several irregular minor ones) each of which has a considerable quantity of secondary tissue all round it, so that a portion of the secondary wood is growing in towards the actual centre of the stem as a whole—a very anomalous state of affairs.

In the more complex Continental type of _Medullosa_ there are _very_ large numbers of steles. In the one figured from the Continent in fig. 51 but a few are represented. There is a large outer double-ring stele, with secondary wood on both sides of it, and within these a number of small steles, all scattered through the ground tissue, and each surrounded by secondary wood. In actual specimens the number of these central steles is much greater than that indicated in the diagram.

No plant exists to-day which has such an arrangement of its vascular cylinder. It almost appears as though at the early period, when the Medulloseæ flourished, steles were experimenting in various directions. Such types as are illustrated in figs. 50 and 51 are obviously wasteful (for secondary wood developing towards the centre of a stem is bound to finally meet), and complex, but apparently inefficient, which may partly account for the fact that this type of structure has not survived to the present, though simpler and equally ancient types have done so.

[Illustration: Fig. 51.—Continental _Medullosa_, showing R, outer double-ring stele with secondary wood all round it; S, inner stellate steles, also surrounded in each case by secondary tissue]

Further details of the anatomy of fossils will be mentioned when we come to consider the individual families; those now illustrated suffice to show that in the Coal Measures very different arrangements of steles were to be found, as well as those which were similar to those existing now. The significance of these differences will become apparent when their relation to the other characters of the plants is considered.

The fructifications, always the most important parts of the plant, offer a wide field, and the divergence between the commoner palæozoic and recent types seems at first to be very great. Indeed, when palæozoic reproductive bodies have to be described, it is often necessary to use the common descriptive terms in an altered and wider sense.

Among the plants of to-day there are many varieties of the simple single-celled reproductive masses which are called _spores_, and which are usually formed in large numbers inside a spore case or sporangium. Among the higher plants _seeds_ are also known in endless variety, all of which, compared with spores, are very complex, for they are many-celled structures, consisting essentially of an embryo or young plant enclosed in various protective coats. The distinction between the two is sharp and well defined, and for the student of living plants there exists no difficulty in separating and describing seeds and spores.

But when we look back through the past eras to palæozoic plants the subject is not so easy, and the two main types of potentially reproductive masses are not sharply distinct. The seed, as we know it among recent plants, and as it is generally defined, had not fully evolved; while the spores were of great variety and had evolved in several directions, some of which seem to have been intermediate stages between simple spores and true seeds. These seedlike spores served to reproduce the plants of the period, but their type has since died out and left but two main methods among living plants, namely the essentially simple spores, the very simplicity of whose organization gives them a secure position, and the complex seeds with their infinite variety of methods for protecting and scattering the young embryos they contain.

Among the Coal Measure fossils we can pick up some of the early stages in the evolution of the seed from the spore, or at least we can examine intermediate stages between them which give some idea of the possible course of events. Hence, though the differences from our modern reproductive structures are so noticeable a feature of the palæozoic ones, it will be seen that they are really such differences as exist between the members at the two ends of a series, not such as exist between unrelated objects.

Very few types can be mentioned here, and to make their relations clear a short series of diagrams with explanations will be found more helpful than a detailed account of the structures.

[Illustration: Fig. 52.—Spores

Each spore a single cell which develops with three others in tetrads (groups of four). Very numerous tetrads enclosed in a spore case or sporangium which develops on a leaflike segment called the sporophyll. Each spore germinates independently of the others after being scattered, all being of the same size. Common in fossils and living Pteridophytes.]

[Illustration: Fig. 53.—Spores

Each a single cell like the preceding, but here only one tetrad in a sporangium ripens, so that each contains only four spores. Compared with the preceding types these spores are very large. Otherwise details similar to above. Some fossils have such sporangia with eight spores, or some other small number; living Selaginellas have four. In the same cone sporangia with small spores are developed and give rise to the male organs.]

[Illustration: Fig. 54.—“Spores” of Seedlike Structure

Out of a tetrad in each sporangium only one spore ripens, S in figure, the others, _s_, abort. The wall of the sporangium, _w_, is more massive than in the preceding cases, and from the sporophyll, flaps, _sp f_, grow up on each side and enclose and protect the sporangium. The one big spore appears to germinate inside these protective coats, and not to be scattered separately from them. Only found in fossils, one of the methods of reproduction in _Lepidodendron_. Other sporangia with small spores were developed which gave rise to the male organs.]

[Illustration: Fig. 55.—“Seed”

In appearance this is like a seed, but differs from a true seed in having no embryo, and is like the preceding structure in having a very large spore, S, though there is no trace of the three aborting ones. The spore develops in a special mass of tissue known as the nucellus, _n_, which partly corresponds to the sporangium wall of the previous types. In it a cavity, _p c_, the pollen chamber, receives the pollen grains which enter at the apex of the “seed”. There is a complex coat, C, which stands round the nucellus but is not joined to it, leaving the space _l_ between them. Only in fossils; _Trigonocarpus_ (see p. 122) is similarly organized. Small spores in fern-like sporangia, called pollen grains.]

[Illustration: Fig. 56.—“Seed”

Very similarly organized to the above, but the coat is joined to the nucellus about two-thirds of its extent, and up to the level _l_. In the pollen chamber, _p c_, a cone of nucellar tissue projects, and the upper part of the coat is fluted, but these complexities are not of primary importance. The large spore S germinated and was fertilized within the “seed”, but apparently produced no embryo before it ripened. Small “spores” in fern-like sporangia form the pollen grains. Only in fossils, _e.g._ Lagenostoma. (See p. 119.)]

[Illustration: Fig. 57.—Seed

Essentially similar to the preceding, except in the possession of an embryo _e_, which is, however, small in comparison with the endosperm which fills the spore S. The whole organization is simpler than in the fossil _Lagenostoma_, but the coat is fused to the nucellus further up (see _l_). Small “spores” form the pollen grains. Living and fossil type, Cycads and Ginkgo.]

[Illustration: Fig. 58.—Seed

In the ripe seed the large embryo _e_ practically fills up all the space within the two seed coats _c^1_ and _c^2_; endosperm, pollen chamber, &c., have been eliminated, and the young ovule is very simple and small as a result of the protection and active service of the carpels in which it is enclosed. Small “spores” form the pollen grains. Typical of living Dicotyledons.]

These few illustrations represent only the main divisions of an army of structures with an almost unimaginable wealth of variety which must be left out of consideration.

For the structures illustrated in figs. 54, 55, and 56 we have no name, for their possible existence was not conceived of when our terminology was invented, and no one has yet christened them anew with distinct names. They are evidently too complex in organization and too similar to seeds in several ways to be called spores, yet they lack the essential element in a seed, namely, an embryo. The term “ovule” (usually given to the young seed which has not yet developed an embryo) does not fit them any better, for their tissues are ripened and hard, and they were of large size and apparently fully grown and mature.

For the present a name is not essential; the one thing that is important is to recognize their intermediate character and the light they throw on the possible evolution of modern seeds.

A further point of great interest is the manner in which these “seeds” were borne on the plant. To-day seeds are always developed (with the exception of Cycas) in cones or flowers, or at least special inflorescences. But the “seed” of _Lagenostoma_ (fig. 56), as well as a number of others in the group it represents, were not borne on a special structure, but directly on the green foliage leaves. They were in this on a level with the simple sporangia of ferns which appear on the backs of the fronds, a fact which is of great significance both for our views on the evolution of seeds as such, and for the bearing it has on the relationships of the various groups of allied plants. This will be referred to subsequently (Chapter XI), and is mentioned now only as an example of the difference between some of the characters of early fossils and those of the present day.

It is true that botanists have long recognized the organ which bears seeds as a modified leaf. The carpels of all the higher plants are looked on as _homologous_ with leaves, although they do not appear to be like them externally. Sometimes among living plants curious diseases cause the carpels to become foliar, and when this happens the diseased carpel reverts more or less to the supposed ancestral leaf-like condition. It is only among the ancient (but recently discovered) fossils, however, that seeds are known to be borne normally on foliage leaves.

From Mesozoic plants we shall learn new conceptions about flowers and reproductive inflorescences in general, but these must be deferred to the consideration of the family as a whole (Chapter XIII).

Enough has been illustrated to show that though the individual cells, the bricks, so to speak, of plant construction, were so similar in the past and present, yet the organs built up by them have been continually varying, as a child builds increasingly ambitious palaces with the same set of bricks.

CHAPTER VIII PAST HISTORIES OF PLANT FAMILIES I. Flowering Plants, Angiosperms

In comparison with the other groups of plants the flowering families are of recent origin, yet in the sense in which the word is usually used they are ancient indeed, and the earliest records of them must date at least to periods hundreds of thousands of years ago.

Through all the Tertiary period (see p. 34) there were numerous flowering plants, and there is evidence that many families of both Monocotyledons and Dicotyledons existed in the Upper Cretaceous times. Further back than this we have little reliable testimony, for the few specimens of so-called flowering plants from the Lower Mesozoic are for the most part of a doubtful nature.

The flowering plants seem to stand much isolated from the rest of the plant world; there is no _direct_ evidence of connection between their oldest representatives and any of the more primitive families. So far as our actual knowledge goes, they might have sprung into being at the middle of the Mesozoic period quite independently of the other plants then living; though there are not wanting elaborate and almost convincing theories of their connection with more than one group of their predecessors (see p. 108).

It is a peculiarly unfortunate fact that although the rocks of the Cretaceous and Tertiary are so much less ancient than those of the Coal Measures, they have preserved for us far less well the plants which were living when they were formed. Hitherto no one has found in Mesozoic strata masses of exquisitely mineralized Angiosperm fragments[8] like those found in the Coal Measures, which tell us so much about the more ancient plants. Cases are known of more or less isolated fragments with their microscopical tissues mineralized. For example, there are some palms and ferns from South America which show their anatomical structure very clearly preserved in silica, and which seem to resemble closely the living species of their genera. The bulk of the plants preserved from these periods are found in the form of casts or impressions (see p. 10), which, as has been pointed out already, are much less satisfactory to deal with, and give much less reliable results than specimens which have also their internal structure petrified. The quantity of material, however, is great, and impressions of single leaves innumerable, and of specimens of leaves attached to stems, and even of flowers and fruits, are to be found in the later beds of rock. These are generally clearly recognizable as belonging to one or other of the living families of flowering plants. Leaf impressions are by far the most frequent, and our knowledge of the Tertiary flora is principally derived from a study of them. Their outline and their veins are generally preserved, often also their petioles and some indication of the thickness and character of the fleshy part of the leaf. From the outline and veins alone an expert is generally able to determine the species to which the plant belongs, though it is not always quite safe to trust to these determinations or to draw wide-reaching conclusions from them.

In fig. 59 is shown a photograph of the impression of a Tertiary leaf, which illustrates the condition of an average good specimen from rocks of the period. Its shape and the character of the veins are sufficient to mark it out immediately as belonging to the Dicotyledonous group of the flowering plants.

Seeds and fruits are also to be found; and in some very finely preserved specimens from Japan stamens from a flower and delicate seeds are seen clearly impressed on the light stone. In fig. 60 is illustrated a couple of such seeds, which show not only their wings but also the small antennæ-like stigmas. Specimens so perfectly preserved are practically as good as herbarium material of recent plants, and in this way the externals of the Tertiary plants are pretty well known to us.

[Illustration: Fig. 59.—Dicotyledonous Leaf Impression from Tertiary Rocks]

[Illustration: Fig. 60.—Seeds from Japanese Tertiary Rocks; at _a_ are seen the two stigmas still preserved]

A problem which has long been discussed, and which has aroused much interest, is the relative antiquity of the Monocotyledonous and the Dicotyledonous branches of the flowering plants. A peculiar fascination seems to hang over this still unsolved riddle, and a battle of flowers may be said to rage between the lily and the rose for priority. Recent work has thrown no decisive light on the question, but it has undoubtedly demolished the old view which supposed that the Monocotyledons (the lily group) appeared at a far earlier date upon this earth than the Dicotyledons. The old writers based their contention on incorrectly determined fossils. For instance, seeds from the Palæozoic rocks were described as Monocotyledons because of the three or six ribs which were so characteristic of their shell; we know now that these seeds (_Trigonocarpus_) belong to a family already mentioned in another connection (p. 72), the Medulloseæ (see p. 122), the affinity of which lies between the cycads and the ferns. Leaves of _Cordaites_, again, which are broad and long with well-marked parallel veins, were described as those of a Monocotyledonous plant like the Yucca of to-day; but we now know them to belong to a family of true Gymnosperms possibly distantly related to _Taxus_ (the Yew tree).

Recent work, which has carefully sifted the fossil evidence, can only say that no true Monocotyledons have yet been found below the Lower Cretaceous rocks, and that at that period we see also the sudden inrush of Dicotyledons. Hence, so far as palæontology can show, the two parallel groups of the flowering plants arose about the same time. It is of interest to note, however, that the only petrifaction of a flower known from any part of the world is an ovary which seems to be that of one of the Liliaceæ. In the same nodules, however, there are several specimens of Dicotyledonous woods, so that it does not throw any light on the question of priority.

With the evidence derived from the comparative study of the anatomy of recent flowering plants we cannot concern ourselves here, beyond noting that the results weigh in favour of the Dicotyledons as being the more primitive, though not necessarily developed much earlier in point of time. Until very much more is discovered than is yet known of the origin of the flowering plants as a whole, it is impossible to come to a more definite conclusion about this much-discussed subject.

Let us now attempt to picture the vegetable communities since the appearance of the flowering plants. The facts which form the bases of the following conceptions have been gathered from many lands by numerous workers in the field of fossil botany, from scattered plant remains such as have been described.

When the flowering plants were heralded in they appeared in large numbers, and already by the Cretaceous period there were very many different species. Of these a number seem to belong to genera which are still living, and many of them are extremely like living species. It would be wearisome and of little value to give a list of all the recorded species from this period, but a few of the commoner ones may be mentioned to illustrate the nature of the plants then flourishing.

Several species of _Quercus_ (the Oak) appeared early, particularly _Quercus Ilex_; leaves of the _Juglandaceæ_ (Walnut family) were very common, and among the Tertiary fossils appear its fruits. Both _Populus_ (the Poplar) and _Salix_ (the Willow) date from the early rocks, while _Ficus_ (the Fig) was very common, and _Casuarina_ (the Switch Plant) seems to have been widely spread. Magnolias also were common, and it appears that _Platanus_ (the Plane) and _Eucalyptus_ coexisted with them.

It will be immediately recognized that the above plants have all living representatives, either wild or cultivated, growing in this country at the present day, so that they are more or less familiar objects, and there appears to have been no striking difference between the early flowering plants and those of the present day. Between the ancient Lycopods, for example, and those now living the differences are very noteworthy; but the earliest of the known flowering plants seem to have been essentially like those now flourishing. It must be remembered in this connection that the existing flowering plants are immensely nearer in point of time to their origin than are the existing Lycopods, and that when such æons have passed as divide the present from the Palæozoic, the flowering plants of the future may have dwindled to a subordinate position corresponding to that held by the Lycopods now.

A noticeable character of the early flowering-plant flora, when taken as a whole, is the relatively large proportion of plants in it which belong to the family _Amentiferæ_ (oaks, willows, poplars, &c.). This is supposed by some to indicate that the family is one of the most primitive stocks of the Angiosperms. This view, however, hardly bears very close scrutiny, because it derives its main support from the large numbers of the Amentiferæ as compared with other groups. Now, the Amentiferæ were (and are) largely woody resistant plants, whose very nature would render them more liable to be preserved as impressions than delicate trees or herbs, which would more readily decay and leave no trace. Similarly based on uncertain evidence is the surmise that the group of flowers classed as _Gamopetalæ_ (flowers with petals joined up in a tube, like convolvulus) did not flourish in early times, but are the higher and later development of the flower type. Now, _Viburnum_ (allied to the honeysuckle) belongs to this group, and it is found right down in the Cretaceous, and _Sambucus_ (Elder, of the same family) is known in the early Tertiary. These two plants are woody shrubs or small trees, while many others of the family are herbs, and it is noteworthy that it is just these woody, resistant forms which are preserved as fossils; their presence demonstrates the antiquity of the group as a whole, and the absence of other members of it may be reasonably attributed to accidents of preservation. In the Tertiary also we get a member of the heath family, viz. _Andromeda_, and another tube-flower, _Bignonia_, as well as several more _woody_ gamopetalous flowers.

Hence it is wise to be very cautious about drawing any important conclusions from the relative numbers of the different species, or the absence of any type of plant from the lists of those as yet known from the Cretaceous. When quantities of structurally preserved material can be examined containing the flowering plants in petrifactions, then it will be possible to speak with some security of the nature of the Mesozoic flora as a whole.

The positive evidence which is already accumulated, however, is of great value, and from it certain deductions may be safely made. Specimens of Cretaceous plants from various parts of the world seem to indicate that there was a very striking uniformity in the flora of that period all over the globe. In America and in Central Europe, for example, the same types of plants were growing. We shall see that, as time advanced, the various types became separated out, dying away in different places, until each great continent and division of land had a special set of plants of its own. At the commencement of the reign of flowering plants, however, they seem to have lived together in the way we are told the beasts first lived in the garden of Eden.

At the beginning of the Tertiary period there were still many tropical forms, such as Palms, Cycads, _Nipa_, various _Artocarpaceæ_, _Lauraceæ_, _Araliaceæ_, and others, growing side by side with such temperate forms as _Quercus_, _Alnus_, _Betula_, _Populus_, _Viburnum_, and others of the same kind. Before the middle of the Tertiary was reached the last Cycads died in what is now known as Europe; and soon after the middle Tertiary all the tropical types died out of this zone.

At the same time those plants whose leaves appear to have fallen at the end of the warm season began to become common, which is taken as an indication of a climatic influence at work. Some writers consider that in the Cretaceous times there was no cold season, and therefore no regular period of leaf fall, but as the climate became temperate the deciduous trees increased in numbers; yet the Gymnospermic and Angiospermic woods which are found with petrified structure show well-marked annual rings and seem to contradict this view.

Toward the end of the Tertiary times there were practically no more tropical forms in the European flora, though there still remained a number of plants which are now found either only in America or only in Asia.

The Glacial epoch at the close of the Tertiary appears to have driven all the plants before it, and afterwards, when its glaciers retreated, shrinking up to the North and up the sides of the high mountains, the plant species that returned to take possession of the land in the Quaternary or present period were those which are still inhabiting it, and the floras of the tropics, Asia, and America were no longer mixed with that of Europe.[9]

CHAPTER IX PAST HISTORIES OF PLANT FAMILIES II. Higher Gymnosperms

The more recent history of the higher Gymnosperms, in the Upper Cretaceous and Tertiary periods, much resembles that of the flowering plants as sketched in the previous chapter. Many of the genera appear to have been those still living, and some of the species even may have come very close to or have been identical with those of to-day. The forms now characteristic of the different continents were growing together, and appear to have been widely distributed over the globe. For example, _Sequoia_ and _Taxodium_, two types now characteristic of America, and _Glyptostrobus_, at present found in Asia, were still growing with the other European types in Europe so late as middle Tertiary times.

As in the case of the Angiosperms, the fossils we have of Cretaceous and Tertiary Gymnosperms are nearly all impressions and casts, though some more or less isolated stems have their structure preserved. Hence our knowledge of these later Gymnosperms is far from complete. From the older rocks, however, we have both impressions and microscopically preserved material, and are more fully acquainted with them than with those which lived nearer our own time. Hard, resistant leaves, which are so characteristic of most of the living genera of Gymnosperms, seem to have been also developed in the past members of the group, and these tend to leave clear impressions in the rocks, so that we have reliable data for reconstructing the external appearance of the fossil forms from the Palæozoic period.

The resinous character of Gymnosperm wood probably greatly assisted its preservation, and fragments of it are very common in rocks of all ages, generally preserved in silica so as to show microscopic structure. The isolated wood of Gymnosperms, however, is not very instructive, for from the wood alone (and usually it is just fragments of the secondary wood which are preserved) but little of either physiological or evolutional value can be learned. When twigs with primary tissues and bark and leaves attached are preserved, then the specimens are of importance, for their true character can be recognized. Fortunately among the coal balls there are many such fragments, some of which are ac