CHAPTER V.. The Flora of the Early Mesozoic.

THE FLORA OF THE EARLY MESOZOIC.

Great physical changes occurred at the close of the Carboniferous age. The thick beds of sediment that had been accumulating in long lines along the primitive continents had weighed down the earth's crust. Slow subsidence had been proceeding from this cause in the coal-formation period, and at its close vast wrinklings occurred, only surpassed by those of the old Laurentian time. Hence in the Appalachian region of America we have the Carboniferous beds thrown into abrupt folds, their shales converted into hard slates, their sandstones into quartzite and their coals into anthracite, and all this before the deposition of the Triassic Red Sandstones which constitute the earliest deposit of the great succeeding Mesozoic period. In like manner the coal-fields of Wales and elsewhere in western Europe have suffered similar treatment, and apparently at the same time.

This folding is, however, on both sides of the Atlantic limited to a band on the margin of the continents, and to certain interior lines of pressure, while in the middle, as in Ohio and Illinois in America, and in the great interior plains of Europe, the coal-beds are undisturbed and unaltered. In connection with this we have an entire change in the physical character of the deposits, a great elevation of the borders of the continents, and probably a considerable deepening of the seas, leading to the establishment of general geographical conditions which still remain, though they have been temporarily modified by subsequent subsidences and re-elevations.

Along with this a great change was in progress in vegetable and animal life. The flora and fauna of the Palæozoic gradually die out in the Permian and are replaced in the succeeding Trias by those of the Mesozoic time. Throughout the Permian, however, the remains of the coal-formation flora continue to exist, and some forms, as the Calamites, even seem to gain in importance, as do also certain types of coniferous trees. The Triassic, as well as the Permian, was marked by physical disturbances, more especially by great volcanic eruptions discharging vast beds and dykes of lava and layers of volcanic ash and agglomerate. This was the case more especially along the margins of the Atlantic, and probably also on those of the Pacific. The volcanic sheets and dykes associated with the Red Sandstones of Nova Scotia, Connecticut, and New Jersey are evidences of this.

At the close of the Permian and beginning of the Trias, in the midst of this transition time of physical disturbance, appear the great reptilian forms characteristic of the age of reptiles, and the earliest precursors of the mammals, and at this time the old Carboniferous forms of plants finally pass away, to be replaced by a flora scarcely more advanced, though different, and consisting of pines, cycads, and ferns, with gigantic equiseta, which are the successors of the genus Calamites, a genus which still survives in the early Trias. Of these groups the conifers, the ferns, and the equiseta are already familiar to us, and, in so far as they are concerned, a botanist who had studied the flora of the Carboniferous would have found himself at home in the succeeding period. The cycads are a new introduction. The whole, however, come within the limits of the cryptogams and the gymnosperms, so that here we have no advance.

Fontaine's "Early Mesozoic Flora of Virginia" gives a very good summary of this flora in America.

As we ascend, however, in the Mesozoic, we find new and higher types. Even within the Jurassic epoch, the next in succession to the Trias, there are clear indications of the presence of the endogens, in species allied to the screw-pines and grasses; and the palms appear a little later, while a few exogenous trees have left their remains in the Lower Cretaceous, and in the Middle and Upper Cretaceous these higher plants come in abundantly and in generic forms still extant, so that the dawn of the modern flora belongs to the Middle and Upper Cretaceous. It will thus be convenient to confine ourselves in this chapter to the flora of the earlier Mesozoic.

Passing over for the present the cryptogamous plants already familiar in older deposits, we may notice the new features of gymnospermous and phænogamous life, as they present themselves in this earlier part of the great reptilian age, and as they extended themselves with remarkable uniformity in this period over all parts of the world. For it is a remarkable fact that, if we place together in our collections fossil plants of this period from Australia, India, China, Siberia, Europe, or even from Greenland, we find wonderfully little difference in their aspect. This uniformity we have already seen prevailed in the Palæozoic flora; and it is perhaps equally marked in that of the Mesozoic. Still we must bear in mind that some of the plants of these periods, as the ferns and pines, for example, are still world-wide in their distribution; but this does not apply to others, more especially the cycads (Fig. 65).

The cycads constitute a singular and exceptional type in the modern world, and are limited at present to the warmer climates, though very generally distributed in these, as they occur in Africa, India, Japan, Australia, Mexico, Florida, and the West Indies. In the Mesozoic age, however, they were world-wide in their distribution, and are found as far north as Greenland, though most of the species found in the Cretaceous of that country are of small size/ and may have been of low growth, so that they may have been protected by the snows of winter. The cycads have usually simple or unbranching stems, pinnate leaves borne in a crown at top, and fruits which, though somewhat various in structure and arrangement, are all of the simpler form of gymnospermous type. The stems are exogenous in structure, but with slender wood and thick bark, and barred tissue, or properly as tissue intermediate between this and the disc-bearing fibres of the pines.

Though the cycads have a considerable range of organisation and of fructification, and though some points in reference to the latter might assign them a higher place, on the whole they seem to occupy a lower position than the conifers or the cordaiteæ of the Carboniferous. In the Carboniferous some of the fern-like leaves assigned to the genus Noeggerathia have been shown by Stur and Weiss to have been gymnosperms, probably allied to cycads, of which they may be regarded at least as precursors. Thus the cycadean type does not really constitute an advance in grade of organisation in the Mesozoic, any further than that, in the period now in question, it becomes much more developed in number and variety of forms. But the conifers would seem to have had precedence of it for a long time in the Palæozoic, and it replaces in the Mesozoic the Cordaites, which in many respects excelled it in complexity.

The greater part of the cycads of the Mesozoic age would seem to have had short stems and to have constituted the undergrowth of woods in which conifers attained to greater height. An interesting case of this is the celebrated dirt-bed of the quarries of the Isle of Portland, long ago described by Dean Buckland. In this fossil soil trunks of pines, which must have attained to great height, are interspersed with the short, thick stems of cycads, of the genus named Cycadoidea by Buckland, and which from their appearance are called "fossil birds' nests" by the quarrymen. Some, however, must have attained a considerable height so as to resemble palms.

The cycads, with their simple, thick trunks, usually marked with rhombic scars, and bearing broad spreading crowns of large, elegantly formed pinnate leaves, must have formed a prominent part of the vegetation of the northern hemisphere during the whole of the Mesozoic period. A botanist, had there been such a person at the time, would have found this to be the case everywhere from the equator to Spitzbergen, and probably in the southern hemisphere as well, and this throughout all the long periods from the Early Trias to the Middle Cretaceous. In a paper published in the "Linnæan Transactions" for 1868, Dr. Carruthers enumerates twenty species of British Mesozoic cycads, and the number might now be considerably increased.

The pines present some features of interest. We have already seen their connection with the broad-leaved Cordaites, and in the Permian there are some additional types of broad-leaved coniferæ. In the Mesozoic we have great numbers of beautiful trees, with those elegant fan-shaped leaves characteristic of but one living species, the Salisburia, or gingko-tree of China. It is curious that this tree, though now limited to eastern Asia, will grow, though it rarely fruits, in most parts of temperate Europe, and in America as far north as Montreal, and that in the Mesozoic period it occupied all these regions, and even Siberia and Greenland, and with many and diversified species (Fig. 66).

Salisburia belongs to the yews, but an equally curious fact applies to the cypresses. The genus Sequoia, limited at present to two species, both Californian, and one of them the so-called "big tree," celebrated for the gigantic size to which it attains, is represented by species found as far back at least as the Lower Cretaceous, and in every part of the northern hemisphere. It seems to have thriven in all these regions throughout the Mesozoic and early Kainozoic, and then to have disappeared, leaving only a small remnant to represent it in modern days. A number of species have been described from the Mesozoic and Tertiary, all of them closely related to those now existing (Fig. 67).

In the Eocene of Australia.

The following notice of these trees is for the most part translated, with some modifications and abridgment, from a paper read by the late Prof. Heer before the Botanical Section of the Swiss Natural History Society:

The name itself deserves consideration. It is that of an Indian of the Cherokee tribe, Sequo Yah, who invented an alphabet without any aid from the outside world of culture, and taught it to his tribe by writing it upon leaves. This came into general use among the Cherokees, before the white man had any knowledge of it; and afterward, in 1828, a periodical was published in this character by the missionaries. Sequo Yah was banished from his home in Alabama, with the rest of his tribe, and settled in New Mexico, where he died in 1843.

When Endlicher was preparing his synopsis of the conifers, in 1846, and had established a number of new genera, Dr. Jacbon Tschudi, then living with Endlicher, brought before his notice this remarkable man, and asked him to dedicate this red-wooded tree to the memory of a literary genius so conspicuous among the red men of America. Endlicher consented to do so, and only endeavored to make the name pronounceable by changing two of its letters.

Endlicher founded the genus on the redwood of the Americans, Taxodium sempervirens of Lamb; and named the species Sequoia sempervirens. These trees form large forests in California, which extend along the coast as far as Oregon. Trees are there met with of 300 feet in height and 20 feet in diameter. The seeds have been brought to Europe a number of years ago, and we already see in upper Italy and around the Lake of Geneva, and in England, high trees; but, on the other hand, they have not proved successful around Zurich.

In 1852, a second species of Sequoia was discovered in California, which, under the name of big tree, soon attained a considerable celebrity. Lindley described it, in 1853, as Wellingtonia gigantea; and, in the following year, Decaisne and Torrey proved that it belonged to Sequoia, and that it accordingly should be called Sequoia gigantea.

While the Sequoia sempervirens, in spite of the destructiveness of the American lumbermen, still forms large forests along the coast, the Sequoia gigantea is confined to the isolated clumps which are met with inland at a height of 5,000 to 7,000 feet above sea-level, and are much sought after by tourists as one of the wonders of the country. Reports came to Europe concerning the largest of them which were quite fabulous, but we have received accurate accounts of them from Prof. Whitney. The tallest tree measured by him has a height of 325 feet, and in the case of one of the trees the number of the rings of growth indicated an age of about 1,300 years. It had a girth of 50 to 60 feet.

We know only two living species of Sequoia, both of which are confined to California. The one (S. sempervirens) is clothed with erect leaves, arranged in two rows, very much like our yew-tree, and bears small, round cones; the other (S. gigantea) has smaller leaves, set closely against the branches, giving the tree more the appearance of the cypress. The cones are egg-shaped, and much larger. These two types are therefore sharply defined.

Both of these trees have an interesting history. If we go back into the Tertiary, this same genus meets us with a long array of species. Two of these species correspond to those living at present: the S. Langsdorfii to the S. sempervirens, and the S. Couttsiæ to the S. gigantea. But, while the living species are confined to California, in the Tertiary they are spread over several quarters of the globe.

S. Couttsiæ has leaves like S. gigantea, and cones like those of 8. sempervirens.

Let us first consider the Sequoia Langsdorfii. This was first discovered in the lignite of Wetterau, and was described as Taxites langsdorfii. Heer found it in the upper Rhone district, and there lay beside the twigs the remains of a cone, which showed that the Taxites Langsdorfii of Brongniart belonged to the Californian genus Sequoia established by Endlicher. He afterward found much better preserved cones, together with seeds, along with the plants of east Greenland, which fully confirmed the determination. At Atanekerdluk in Greenland (about 70° north latitude) this tree is very common. The leaves, and also the flowers and numerous cones, leave no doubt that it stands very near to the modern redwood. It differs from it, however, in having a much larger number of scales in the cone. The tree is also found in Spitzbergen at nearly 78° north latitude, where Nordenskiöld has collected, at Cape Lyell, wonderfully preserved branches. From this high latitude the species can be followed down through the whole of Europe as far as the middle of Italy (at Senegaglia, Gulf of Spezia). In Asia, also,, we can follow it to the steppes of Kirghisen, to Possiet, and to the coast of the Sea of Japan, and across to Alaska and Sitka. It is recognized by Mr. Starkie Gardner as one of the species found in the Eocene of Mull in the Hebrides. It is thus known in Europe, Asia, and America, from 43° to 78° north latitude, while its most nearly related living species, perhaps even descended from it, is now confined to California.

It is Fareites Campbelli of Forbes.

With this S. Langsdorfii, three other Tertiary species are nearly related (S. brevifolia, Hr., S. disticha, Hr., and S. Nordenskiöldi, Hr.). These have been met with in Greenland and Spitzbergen, and one of them has lately been found in the United States. Three other species, in addition to these, have been described by Lesquereux, which appear to belong to the group of the S. Langsdorfii, viz., S. longifolia, Lesq., S. angustifolia, and S. acuminata, Lesq. Several species also occur in the Cretaceous and Eocene of Canada.

These species thus answer to the living Sequoia sempervirens; but we can also point to Tertiary representatives of the S. gigantea. Their leases are stiff and sharp-pointed, are thinly set round the branches, and lie forward in the same way: the egg-shaped cones are in some cases similar.

There are, however, in the early Tertiary six species, which fill up the gap between S. sempervirens and S. gigantea. They are the S. Couttsiæ, S. affinis, Lesq., S. imbricata, Hr., S. sibirica, Hr., S. Heerii, Lesq., and S. biformis, Lesq. Of these, S. Couttsiæ, Hr., is the most common and most important species. It has short leaves, lying along the branch, like S. gigantea, and small, round cones, like S. Langsdorfii and sempervirens. Bovey Tracey in Devonshire has afforded splendid specimens of cones, seeds, and twigs, which have been described in the "Philosophical Transactions." More lately, Count Saporta has described specimens of cones and twigs from Armissan. Specimens of this species have also been found in the older Tertiary of Greenland, so that it must have had a wide range. It is very like to the American S. affinis, Lesq.

In the Tertiary there have been already found fourteen well-marked species, which thus include representatives of the two living types, S. sempervirens and S. gigantea.

We can follow this genus still further back. If we go back to the Cretaceous age, we find ten species, of which five occur in the Urgon of the Lower Cretaceous, two in the Middle, and three in the Upper Cretaceous. Among these, the Lower Cretaceous exhibits the two types of the Sequoia sempervirens and S. gigantea. To the former the S. Smithiana answers, and to the latter, the Reichenbachii, Gein. The S. Smithiana stands indeed uncommonly near the S. Langsdorfii, both in the appearance of the leaves on the twigs and in the shape of the cones. These are, however, smaller, and the leaves do not become narrower toward the base. The S. pectina, Hr., of the Upper Cretaceous, has its leaves arranged in two rows, and presents a similar appearance. The S. Reichenbachii is a type more distinct from those now living and those in the Tertiary. It has indeed stiff, pointed leaves, lying forward, but they are arcuate, and the cones are smaller. This tree has been known for a long time, and it serves in the Cretaceous as a guiding star, which we can follow from the Urgonian of the Lower Cretaceous up to the Cenomanian. It is known in France, Belgium, Bohemia, Saxony, Greenland, and Spitzbergen (also in Canada and the United States). It has been placed in another genus--Geinitzia--but we can recognise, by the help of the cones, that it belongs to Sequoia.

Below this, there is found in Greenland a nearly related species, the S. ambigua, Hr., of which the leaves are shorter and broader, and the cones round and somewhat smaller.

The connecting link between S. Smithiana and Reichenbachii is formed by S. subulata, Hr., and S. rigida, Hr., and three species (S. gracilis, Hr., S. fastigiata and S. Gardneriana, Carr.), with leaves lying closely along the branch, and which come very near to the Tertiary species S. Couttsiæ. We have therefore in the Cretaceous quite an array of species, which fill up the gap between the S. sempervirens and gigantea, and show us that the genus Sequoia had already attained a great development in the Cretaceous. This was still greater in the Tertiary, in which it also reached its maximum of geographical distribution. Into the present world the two extremes of the genus have alone continued; the numerous species forming its main body have fallen out in the Tertiary.

If we look still further back, we find in the Jura a great number of conifers, and, among them, we meet in the genus Pinus with a type which is highly developed, and which still survives; but for Sequoia we have till now looked in vain, so that for the present we can not place the rise of the genus lower than the Urgonian of the Cretaceous, however remarkable we may think it that in that period it should have developed into so many species; and it is still more surprising that two species already make their appearance which approach so near to the living Sequoia sempervirens and S. gigantea.

Altogether, we have become acquainted, up to the present time, with twenty-six species of Sequoia. Fourteen of these species are found in the Arctic zone, and have been described and figured in the "Fossil Flora of the Arctic Regions." Sequoia has been recognised by Ettingshausen even in Australia, but there in the Eocene.

This is, perhaps, the most remarkable record in the whole history of vegetation. The Sequoias are the giants of the conifers, the grandest representatives of the family, and the fact that, after spreading over the whole northern hemisphere and attaining to more than twenty specific forms, their decaying remnant should now be confined to one limited region in western America and to two species constitutes a sad memento of departed greatness. The small remnant of S. gigantea still, however, towers above all competitors, as eminently the "big trees "; but, had they and the allied species failed to escape the Tertiary continental submergences and the disasters of the glacial period, this grand genus would have been to us an extinct type. In like manner the survival of the single gingko of eastern Asia alone enables us to understand that great series of taxine trees with fern-like leaves of which it is the sole representative.

The writer has shown that much of the material of the great lignite beds of the Canadian Northwest consists of wood of Sequoia of both the modern types.

Besides these peculiar and now rare forms, we have in the Mesozoic many others related closely to existing yews, cypresses, pines, and spruces, so that the conifers were probably in greater abundance and variety than they are at this day.

In this period, also, we find the earliest representatives of the endogenous plants. It is true that some plants found in the coal-formation have been doubtfully referred to these, but the earliest certain examples would seem to be some bamboo-like and screw-pine-like plants occurring in the Jurassic rocks. Some of these are, it is true, doubtful forms, but of others there seems to be no question. The modern Pandanus or screw-pine of the tropical regions, which is not a pine, however, but a humble relation of the palms, is a stiffly branching tree, of a candelabra-like form, and with tufts of long leaves on its branches, and nuts or great hard berries for fruit, borne sometimes in large masses, and so protected as to admit of their drifting uninjured on the sea. The stems are supported by masses of aërial roots like those which strengthen the stems of tree-ferns. These structures and habits of growth fit the Pandanus for its especial habitat on the shores of tropical islands, to which its masses of nuts are drifted by the winds and currents, and on whose shores it can establish itself by the aid of its aërial roots.

Some plants referred to the cycads have proved veritable botanical puzzles. One of these, the Williamsonia gigas of the English oölite, originally discovered by my friend Dr. Williamson, and named by him Zamia gigas, a very tall and beautiful species, found in rocks of this age in various parts of Europe, has been claimed by Saporta for the Endogens, as a plant allied to Pandanus. Some other botanists have supposed the flowers and fruits to be parasites on other plants, like the modern Rafflesia of Sumatra, but it is possible that after all it may prove to have been an aberrant cycad.

The tree-palms are not found earlier than the Middle Cretaceous, where we shall notice them in the next chapter. In like manner, though a few Angiosperms occur in rocks believed to be Lower or Lower Middle Cretaceous in Greenland and the northwest territory of Canada, and in Virginia, these are merely precursors of those of the Upper Cretaceous, and are not sufficient to redeem the earlier Cretaceous from being a period of pines and cycads.

On the whole, this early Mesozoic flora, so far as known to us, has a monotonous and mean appearance. It no doubt formed vast forests of tall pines, perhaps resembling the giant Sequoias of California; but they must for the most part have been dark and dismal woods, probably tenanted by few forms of life, for the great reptiles of this age must have preferred the open and sunny coasts, and many of them dwelt in the waters. Still we must not be too sure of this. The berries and nuts of the numerous yews and cycads were capable of affording much food. We know that in this age there were many great herbivorous reptiles, like Iguanodon and Hadrosaurus, some of them fitted by their structure to feed upon the leaves and fruits of trees. There were also several kinds of small herbivorous mammals, and much insect life, and it is likely that few of the inhabitants of the Mesozoic woods have been preserved as fossils. We may yet have much to learn of the inhabitants of these forests of ferns, cycads, and pines. We must not forget in this connection that in the present day there are large islands, like New Zealand, destitute of mammalia, and having a flora comparable with that of the Mesozoic in the northern hemisphere, though more varied. We have also the remarkable example of Australia, with a much richer flora than that of the early Mesozoic, yet inhabited only by non-placental mammals, like those of the Mesozoic.

The principal legacy that the Mesozoic woods have handed down to our time is in some beds of coal, locally important, but of far less extent than those of the Carboniferous period. Still, in America, the Richmond coal-field in Virginia is of this age, and so are the anthracite beds of the Queen Charlotte Islands, on the west coast of Canada, and the coal of Brora in Sutherlandshire. Valuable beds of coal, probably of this age, also exist in China, India, and South Africa; and jet, which is so extensively used for ornament, is principally derived from the carbonised remains of the old Mesozoic pines.

In the next chapter we have to study a revolution in vegetable life most striking and unique, in the advent of the forest-trees of strictly modern types.

NOTE TO CHAPTER V.

I append to this chapter a table showing the plant-bearing series of the Cretaceous and Laramie of North America, from a paper in "Trans. R. S. C," 1885, which see for further details:

(In Descending Order.)

Periods. Floras and sub-floras. References. --------------+---------------------------+----------------------------- Transition Upper Laramie or Porcupine { Platanus beds of Souris Eocene to Hill. Fort Union { River and Calgary. Report Cretaceous. group, U. S. territory. { of Geol. Survey of Canada { for 1879, and Memoir of { 1885. --------------+---------------------------+----------------------------- Middle Laramie or Willow Creek beds. { Lemna and Pistia beds of Lower Laramie or St. { bad lands of 49th parallel, Mary River. { Red Deer River, &c., with Upper { lignites. Report 49th Cretaceous { Parallel and Memoir of { 1885. (Danian and Senonian). Fox Hill series Marine.

Fort Pierre series Marine.

{ Sequoia and Brasenia beds Belly River { of S. Saskatchewan, Belly { River, &c. with lignites. { Memoir of 1885.

Coal measures of Nanaimo, { Memoir of 1883. Many B.C., probably here. { dicotyledons, palms, &c. --------------+---------------------------+----------------------------- Middle Dunvegan series of Peace } Cretaceous River. Dakota group, } Memoir of 1883. Many (Turonian and U. S. Amboy clays, } dicotyledons, cycads, &c. Cenomanian). U. S. }

Mill Creek beds of Rocky { Dicotyledonous leaves, Mountains. { similar to Dakota group of { the U. S. Memoir of 1885. --------------+---------------------------+----------------------------- Lower Suskwa River beds and } Cretaceous Queen Charlotte Island } Cycads, pines, a few (Neocomian. coal series. Intermediate } dicotyledons. Report Geol. &c). beds of Rocky } Survey. Memoir of 1885. Mountains. Potomac } series of Virginia. }

Kootanie series of Rocky { Cycads, pines, and ferns. Mountains. { Memoir of 1885. --------------+---------------------------+------------------------------