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 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.
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 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, 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.
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, the Medulloseæ, 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.