Ancient Plants – Past Histories of Plant Families: Higher Gymnosperms

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 accompanied by fruits and male cones, so that we know much of the Palæozoic Gymnosperms, and find that in some respects they differ widely from those now living.

There is, therefore, much more to be said about the fossil Gymnosperms than about the Angiosperms, both because of the better quality of their preservation and because their history dates back to a very much earlier period than does the Angiospermic record. Indeed, we do not know when the Gymnosperms began; the well-developed and ancient group of Cordaiteæ was flourishing before the Carboniferous period, and must therefore date back to the rocks of which we have no reliable information from this point of view, and the origin of the Gymnosperms must lie in the pre-Carboniferous period.

The group of Gymnosperms includes a number of genera of different types, most of which may be arranged under seven principal families. In a sketch of this nature it is, of course, quite impossible to deal with all the less-important families and genera. Those that will be considered here are the following::


Araucareæ, e.g. Monkey-puzzle

Genera both living and fossil.

Fossil forms undoubted so far back as the Jurassic, and presumably further.

Abietineæ, e.g. Pine and Larch

Genera both living and fossil.

Fossils recognized as far back as the Lower Cretaceous.

Cupresseæ, e.g. Juniper, Cypress

Genera both living and fossil.

Fossils recognized as far back as the Jurassic.

Taxeæ, e.g. Yew

Genera living and fossil.

Fossils recognized as far back as the Cretaceous.


Cordaiteæ, e.g. Cordaites

Fossil only.

Characteristic of Devonian, Carboniferous, and Permian periods.

Poroxyleæ, e.g. Poroxylon

Fossil only.

Characteristic of the Carboniferous and Permian.


Ginkgoaceæ, e.g. Ginkgo

Fossil and living, dating back, apparently with little change, to Palæozoic times.

We must pay the most attention to the two last groups, as they are so important as fossils, and the Cordaiteæ were a very numerous family in Coal Measure times. They had their period of principal development so long ago that it is probable that no direct descendants remain to the present time, though some botanists consider that the Taxeæ are allied to them.

Of the groups still living it is difficult, almost impossible, to say which is the highest, the most evolved type. In the consideration of the Gymnosperm family it is brought home with great emphasis how incomplete and partial our knowledge is as yet. Many hold that the Araucareæ are the most primitive of the higher Gymnosperms. In support of this view the following facts are noted. They have a simple type of fructification, with a single seed on a simple scale, and many scales arranged round an axis to form a cone. In the microscopic structure of their wood they have double rows of bordered pits, a kind of wood cell which comes closer to the old fossil types than does the wood of any of the other living genera. Further than this, wood which is almost indistinguishable from the wood of recent Araucarias is found very far back in the rocks, while their leaves are broad and simple, and attached directly to the stem in a way similar to the leaves of the fossil Cordaiteæ, and very different from the needle leaves on the secondary stems of the Pine family; so that there appears good ground for considering the group an ancient and probably a primitive one.

On the other hand, there are not wanting scientists who consider the Abietineæ the living representatives of the most primitive and ancient stock, though on the whole the evidence seems to indicate more clearly that the Pine-tree group is specialized and highly modified. Their double series of foliage leaves, their complex cones (whose structures are not yet fully understood), and their wood all support the latter view.

Some, again, consider the Taxeæ as a very primitive group, and would place them near the Cordaiteæ, with which they may be related. Their fleshy seeds, growing not in cones but on short special axes, support this view, and it is certainly true that in many ways the large seeds, with their succulent coats and big endosperm, are much like those of the lower Gymnosperms and of several fossil types. Those, however, who hold to the view that the Abietineæ are primitive, see in the Taxeæ the latest and most modified type of Gymnosperm.

It will be seen from this that there is no lack of variety regarding the interpretation of Gymnosperm structures.

The Gymnosperms do not stand in such an isolated position as do the Angiosperms. Whatever the variety of views held about the details of the relative placing of the families within the group, all agree in recognizing the evidence which enables us to trace with confidence the connection between the lower Gymnosperms and the families of ferns. There are many indications of the intimate connection between higher and lower Gymnosperms. Between the series exist what might be described as different degrees of cousinship, and in the lower groups lie unmistakable clues to their connection with more ancient groups in the past which bridge over the gaps between them and the ferns.

For the present, however, let us confine ourselves to the history of the more important Gymnosperms, the discussion of their origin and the groups from which they may have arisen must be postponed until the necessary details about those groups have been mentioned.

To a consideration of the living families of Araucareæ, Abietineæ, Cupresseæ, and Taxeæ we can allow but a short space; their general characters and appearance are likely to be known to the reader, and their details can be studied from living specimens if they are not. For purposes of comparison with the fossils, however, it will be necessary to mention a few of the principal features which are of special importance in discussing phylogeny.

The Araucariaceæ are woody trees which attain a considerable size, with broad-based, large leaves attached directly to the stem. In the leaves are a series of numerous parallel vascular bundles. The wood cells in microscopic section show two rows or more of round bordered pits. The cones are very large, but the male and female are different in size and organization. The female cone is composed of series of simple scales arranged spirally round the axis, and each scale bears a single seed and a small ligule.

The pollen grains from the male cone are caught on the ligule and the pollen tubes enter the micropyle of the ovule, bringing in passive male cells which may develop in large numbers in each grain. The seeds when ripe are stony, and some are provided with a wing from part of the tissue of the scale. In the ripe cones the scales separate from the cone axis.

The Abietineæ are woody trees, some reaching a great height, all with a strong main stem. The leaves are of two kinds: primary ones borne directly attached to the stem (as in first-year shoots of the Larch), and secondary ones borne in tufts of two (in Pine) or a large number (in older branches of Larch) on special short branches, the primary leaves only developing as brown scales closely attached to the stems. Leaves generally very fine and needlelike, and with a central vascular bundle. The wood in microscopic section shows a single row of round bordered pits on the narrow tracheæ.

The female cones are large, male and female differing greatly in size and organization. The female cone, composed of a spiral series of pairs of scales, which often fuse together as the cone ripens. Each upper scale of the pair bears two seeds. The pollen grains from the male cone enter the micropyle of the seed and are caught in the tissue (apex of nucellus) there; the pollen tubes discharge passive male cells, only two of which develop in each grain. The seeds when ripe are stony and provided with a wing from the tissue of the scale on which they were borne.

The Cupresseæ are woody trees reaching no great height, and of a bushy, branching growth. The leaves are attached directly to the main stem, and arrange themselves in alternating pairs of very small leaves, closely pressed to the stem. The wood in microscopic section shows a single row of round bordered pits on the tracheæ.

The cones are small, and the scales forming them arranged in cycles. The female scales bear a varying number of seeds. The pollen grain has two passive male cells. The seeds when ripe are stony, with wings, though in some cases (species of Juniper) the cone scales close up and become fleshy, so that the whole fruit resembles a berry.

The Taxeæ are woody, though not great trees, bushily branched. The leaves are attached spirally all round the stem, but place themselves so as to appear to lie in pairs arranged in one horizontal direction. The wood in microscopic section shows a single row of round bordered pits on the tracheæ.

There are small male cones, but the seeds are not borne on cones, growing instead on special short axes, where there may be several young ovules, but on which usually two seeds ripen. The seeds are big, and have an inner stone and outer fleshy covering. Some have special outer fleshy structures known as “arils”, e.g. the red outer cup round the yew “berry” (which is not a berry at all, but a single unenclosed seed with a fleshy coat).

When we turn to the Cordaiteæ we come to a group of plants which bears distinct relationship to the preceding, but which has a number of individual characters. It is a group of which we should know nothing were it not for the fossils preserved in the Palæozoic rocks; yet, notwithstanding the fact that it flourished so long ago, it is a family of which we know much. At the time of the Coal Measures and the succeeding Permo-carboniferous period, it was of great importance, and, indeed, in some of the French deposits it would seem as though whole layers of coal were composed entirely of its leaves.

Among the fossil remains of this family there are impressions, casts, and true petrifactions, so that we know both its external appearance and the internal anatomy of nearly every part of several species of the genus. For a long time the various fossil remains of the plant were not recognized as belonging to each other and together forming the records of one and the same plant:the broad, long leaves with their parallel veins were looked on as Monocotyledons; the pith casts were thought to be peculiar constricted stems, and were called Sternbergia; while the wood, which was known from its microscopic structure, was called Araucarioxylon:but the careful work of many masters of fossil botany, whose laborious studies we cannot describe in detail here, brought all these fragments together and proved them to belong to Cordaites.


Fig. 61.:Leaf of Cordaites, l, attached by its broad base to a Stem, s

We now know that Cordaites were large trees, with strong upright shafts of wood, to whose branches large simple leaves were attached. The leaves were much bigger than those of any living Gymnosperm, even than those of the Kauri Pine (a member of the Araucariaceæ), and seem in some species to have exceeded 3 ft. in length. The trees branched only at the top of the main shaft, and with their huge sword-like leaves must have differed greatly in appearance from any plant now living. The leaves had many parallel veins, as can be seen in fig. 61, and were attached by a broad base directly to the main stem; thus coming closer to the Araucarias than the other groups of Gymnosperms in their leaf characters.


Fig. 62A.:Microscopic Section of Part of a Leaf of Cordaites

V, Vascular bundle; W, wood of bundle; sh, its sheath; S1, large sclerenchyma mass alternating with bundles; S2 and S3, sclerenchyma caps of bundle; P, soft tissue of leaf.

The internal anatomy is often well preserved, and there is a number of species of leaves whose anatomy is known. As will be expected from the parallel veins, in each section there are many vascular bundles running equidistantly through the tissue. Fig. 62A shows the microscopic details from a well-preserved leaf. In all the species patches of sclerenchyma were developed, and everything indicates that they were tough and well protected against loss of water, even to a greater extent than are most of the leaves of living Gymnosperms.

In the stems the pith was much larger than that in living Gymnosperms (where the wood is generally very solid), and it was hollow in older stems, except for discs of tissue across the cavity. The internal cast from these stems has been described before, and is seen in fig. 63.


Fig. 62B.:Much-magnified Wood Elements from Cordaites Stem seen in longitudinal section, the type known as Araucarioxylon. Note the hexagonal outlines of the bordered pits, which lie in several rows

The wood was formed in closely packed radiating rows by a normal cambium, and the tracheæ so formed had characteristic rows of bordered pits. The wood comes nearer to that of the living Araucarias than any other, and indeed the numerous pieces of fossil wood of this type which are known from all the geological periods are called Araucarioxylon. A double strand goes out from the main mass of wood, which afterwards divides and subdivides to provide the numerous bundles of the leaf.


Fig. 63.:Cast of Hollow Pith of Cordaites, the constrictions corresponding to discs of solid tissue across the cavity

In the case of these fossils we are fortunate enough to have the fructifications, both male and female, in a good state of preservation. As in other Gymnosperms, the male and female cones are separate, but they differed less from each other in their arrangement than do those of any of the living types hitherto mentioned. They can hardly be described as true cones, though they had something of that nature; the seeds seem to be borne on special short stems, round which are also sterile scales. In the seed and the way it is borne perhaps the Cordaiteæ may be compared more nearly with the Taxeæ than with the other groups. A seed, not yet ripe, is shown in slightly diagrammatic form in fig. 64, where the essential details are illustrated. The seeds of this family sometimes reached a considerable size, and had a fleshy layer which was thick in comparison with the stone, and externally comparable with a cherry:though, of course, of very different nature in reality, for Cordaites, like Taxus, is a Gymnosperm, with simple naked seeds, while a cherry is the fruit of an Angiosperm.

In a few words, these are the main characters of the large group of Cordaites, which held the dominant position among Gymnosperms in the Palæozoic era. They have relationships, or perhaps one should say likenesses, to many groups. Their stem- and root-anatomy is similar to the Coniferæ of the present day, the position of the ovules is like that in the Taxaceæ, the male cones in some measure recall those of Ginkgo, the anatomy of their leaves has points which are comparable with those of the Cycads, to which group also the large pith in the stem and the structure of some details in the seeds unite them. Their own specially distinctive characters lie in their crown of huge leaves, and unbranched shaft of stem, the similarity of their male and female inflorescences, and some points in their pollen grains which have not been mentioned. The type is a very complex one, possibly coming near the stock which, having branched out in various directions, gave rise to several of the living families.


Fig. 64.:Representation of Cordaites Seed and its Axis with Scales, slightly diagrammatic, modified from Renault.

A, Axis with s, scales; c, coat of the seed, from which the inner parts have shrunk away; n, nucellus; p.c, pollen chamber containing pollen grains which enter through m.

Plants which come very near to the Cordaiteæ are the Poroxyleæ. Of this group we have unfortunately no remains of fructifications in organic connection, so that its actual position must remain a little doubtful till they are discovered. There seems no doubt that they must have borne seeds.

Still, it has been abundantly demonstrated in recent years that the anatomy of the root, stem, and leaves indicates with considerable exactness the position of any plant, so that, as these are known, we can deduce from them, with a feeling of safety, the position that Poroxylon takes in the natural system. In its anatomy the characters are those of the Cordaiteæ, with certain details which show a more primitive nature and seem to be characteristic of the groups below it in organization.

Poroxylon is not common, and until recently had not been found in the Lower Coal Measures of England. The plants appear to have been much smaller than Cordaites, with delicate stems which bore relatively large simple leaves. The anatomy of the root was that common in Gymnosperms, but the stem had a very large pith, and the leaves were much like those of Cordaites in having parallel veins. An important character in the anatomy of the stem was the presence of what is known as centripetal wood. This must be shortly explained. In all the stems hitherto considered, the first-formed wood cells (protoxylems, see p. 57) developed at the central point of the wood, towards the pith. This is characteristic of all Angiosperms and the higher Gymnosperms (except in a couple of recently investigated Pines), but among the lower plants we find that part of the later wood develops to the inner side of these protoxylem masses. The distinction is shown in fig. 65.


Fig. 65.:A, Normal bundle of higher plant; x, protoxylem on inner side next the pith p, and the older wood w outside it, centrifugal wood. B, Bundle with wood cells c developed on inner side of protoxylem, centripetal wood; the arrow indicates the direction of the center of the stem.

This point is one to which botanists have given much attention, and on which they have laid much weight in considering the affinities of the lower Gymnosperms and the intermediate groups between them and the ferns, which are found among the fossils. In Cordaites this point of connection with the lower types is not seen, but in Poroxylon, which has otherwise a stem anatomy very similar to Cordaites, we find groups of centripetal wood developed inside the protoxylem of primary bundles. For this reason, principally, is Poroxylon of interest at present, as in its stem anatomy it seems to connect the Cordaites type with that of the group below it in general organization.

Ginkgoales.:Reference to p. 44 shows that Ginkgo, the Maidenhair tree, belongs to the Ginkgoales, a group taking equal rank with the large and complex series of the Coniferales. The Ginkgoales of the present day, however, have but one living representative. Ginkgo stands alone, the single living species of its genus, representing a family so different from any other living family that it forms a prime group by itself.

Had the tree not been held sacred in China and Japan, it is probable that it would long since have been extinct, for it is now known only in cultivation. It is indeed a relic from the past which has been fortunately preserved alive for our examination. It belongs to the fossil world, as a belated November rose belongs to the summer.

Because of its beauty and interest the plant is now widely distributed under cultivation, and is available for study almost as freely as the other types of living Gymnosperms already mentioned, so that but a short summary of its more important features is needed here.

Old plants, such as can be seen growing freely in Japan (in Kew Gardens there is also a fine specimen), are very tall handsome woody trees, with noble shafts and many branches. The leaves grow on little side shoots and are the most characteristic external feature of the tree; their living form is illustrated in fig. 66, which shows the typical simple shape as well as the lobed form of the leaf which are to be found, with all intermediate stages, on the same tree. No other plant (save a few ferns, which can generally be distinguished from it without difficulty) has leaves at all like these, so that it is particularly easy to identify the fossil remains, of which there are many.


Fig. 66.:A, Tuft of Ginkgo Leaves, showing their “maidenhair”-like shape. B, Single deeply-divided Leaf to be found on the same tree, usually on young branches.

The wood is compact and fine grained, the rings of secondary tissue being developed from a normal cambium as in the case of the higher Gymnosperms, and the individual tracheæ have round bordered pits. There are small male cones, but the seeds are not borne in cones. They develop on special stalks on which are no scales, but a small mass of tissue at the base of the seed called the “collar”. Usually there are two young ovules, of which often only one ripens to a fleshy seed, though both may mature.


Fig. 67.:Ripe Stage of Ginkgo Seeds attached to their Stalk. c, “Collar” of seed.

The ripe seed reaches the size shown in the diagram, and is orange coloured and very fleshy; within it is a stone encasing the endosperm, which is large, green, and starchy, and contains the embryo with two cotyledons. This embryo is small compared with the endosperm, cf. fig. 57, p. 76, which is somewhat similar to that of Ginkgo in this stage.

Of the microscopic characters of the reproductive organs the most remarkable is the male cell. This is not a passive nucleus, as in the plants hitherto considered, but is an actively swimming cell of some size, provided with a spiral of cilia (hairlike structures) whose movements propel it through the water. In the cavity of the unripe seed these swim towards the female cell, and actively penetrate it. The arrangements of the seed are diagrammatically shown in fig. 68, which should be compared with that of Cycas, fig. 76, with which it has many points in common.


Fig. 68.:Section through Seed of Ginkgo

p.c, Pollen chamber in the nucellus n, which is fused to the coat c to the level l; sc, stony layer in coat; S, the big spore, filled with endosperm tissue (in this case green in colour); e, egg cells, one of which will produce the embryo after fertilization.

The nature of the male cell in Cordaites is not yet known, but there is reason to suspect it may have been actively swimming also. As this is uncertain, however, we may consider Ginkgo the most highly organized plant which has such a primitive feature, a feature which is a bond of union between it and the ferns, and which, when it was discovered about a dozen years ago, caused a considerable sensation in the botanical world.

To turn now to the fossil records of this family. Leaf impressions of Ginkgo are found in rocks of nearly all ages back even to the Upper Palæozoic. They show a considerable variety of form, and it is certain that they do not all belong to the same species as the living plant, but probably they are closely allied. Fig. 69 shows a typical impression from the Lower Mesozoic rocks. In this specimen, the cells of the epidermis were fortunately sufficiently well preserved to be seen with the microscope, and there is a distinct difference in the size and shape of the cells of living and fossil species, see fig. 70; but this difference is slight as compared with the great similarity of form and appearance, as can be seen on comparing figs. 69 and 66, B, so that the fossil is at the most a different species of the genus Ginkgo. Among the fossil leaves there is greater variety than among the living ones, and some which are very deeply lobed so as to form a divided palm-like leaf go by different names, e.g. Baiera, but they are supposed to belong to the same family. Fossil seeds and male cones are also known as impressions, and are found far back in the Mesozoic rocks. From the fossil impressions it is certain that Ginkgo and plants closely allied to it were very widespread in the past, as they are found all over Europe as well as the other continents. Particularly in the Lower Mesozoic rocks Ginkgo seems to have been a world-wide type growing in great abundance.


Fig. 69.:Leaf Impression of Ginkgo from Mesozoic Rocks of Scotland


Fig. 70.:Showing Epidermis with Stomates from the lower side of the Leaf seen in fig. 69

e, Epidermis cells; s, stomates; v, long cells of epidermis lying over the veins.

In the Palæozoic the records are not so undoubted, but there is strong evidence which leads us to suppose that if the genus now living were not then extant, at least other closely related genera were, and there seems to be good grounds for supposing that Ginkgo and Cordaites may have both arisen from some ancient common stock.