VEGETATION OF THE LAURENTIAN AND EARLY PALÆOZOIC--QUESTIONS AS TO ALGÆ.
Oldest of all the formations known to geologists, and representing perhaps the earliest rocks produced after our earth had ceased to be a molten mass, are the hard, crystalline, and much-contorted rocks named by the late Sir W. E. Logan Laurentian, and which are largely developed in the northern parts of North America and Europe, and in many other regions. So numerous and extensive, indeed, are the exposures of these rocks, that we have good reason to believe that they underlie all the other formations of our continents, and are even world-wide in their distribution. In the lower part of this great system of rocks which, in some places at least, is thirty thousand feet in thickness, we find no traces of the existence of any living thing on the earth. But, in the middle portion of the Laurentian, rocks are found which indicate that there were already land and water, and that the waters and possibly the land were already tenanted by living beings. The great beds of limestone which exist in this part of the system furnish one indication of this. In the later geological formations the limestones are mostly organic--that is, they consist of accumulated remains of shells, corals, and other hard parts of marine animals, which are composed of calcium carbonate, which the animals obtain directly from their food, and indirectly from the calcareous matter dissolved in the sea-water. In like manner great beds of iron-ore exist in the Laurentian; but in later formations the determining cause of the accumulation of such beds is the partial deoxidation and solution of the peroxide of iron by the agency of organic matter. Besides this, certain forms known as Eozoon Canadense have been recognised in the Laurentian limestones, which indicate the presence at least of one of the lower types of marine animals. Where animal life is, we may fairly infer the existence of vegetable life as well, since the plant is the only producer of food for the animal. But we are not left merely to this inference. Great quantities of carbon or charcoal in the form of the substance known as graphite or plumbago exist in the Laurentian. Now, in more recent formations we have deposits of coal and bituminous matter, and we know that these have arisen from the accumulation and slow putrefaction of masses of vegetable matter. Further, in places where igneous action has affected the beds, we find that ordinary coal has been changed into anthracite and graphite, that bituminous shales have been converted into graphitic shales, and that cracks filled with soft bituminous matter have ultimately become changed into veins of graphite. When, therefore, we find in the Laurentian thick beds of graphite and beds of limestone charged with detached grains and crystals of this substance, and graphitic gneisses and schists and veins of graphite traversing the beds, we recognise the same phenomena that are apparent in later formations containing vegetable débris.
The carbon thus occurring in the Laurentian is not to be regarded as exceptional or rare, but is widely distributed and of large amount. In Canada more especially the deposits are very considerable.
The graphite of the Laurentian of Canada occurs both in beds and in veins, and in such a manner as to show that its origin and deposition are contemporaneous with those of the containing rock. Sir William Logan states that "the deposits of plumbago generally occur in the limestones or in their immediate vicinity, and granular varieties of the rock often contain large crystalline plates of plumbago. At other times this mineral is so finely disseminated as to give a bluish-grey colour to the limestone, and the distribution of bands thus coloured seems to mark the stratification of the rock." He further states: "The plumbago is not confined to the limestones; large crystalline scales of it are occasionally disseminated in pyroxene rock, and sometimes in quartzite and in feldspathic rocks, or even in magnetic oxide of iron." In addition to these bedded forms, there are also true veins in which graphite occurs associated with calcite, quartz, orthoclase, or pyroxene, and either in disseminated scales, in detached masses, or in bands or layers "separated from each other and from the wall-rock by feldspar, pyroxene, and quartz." Dr. Hunt also mentions the occurrence of finely granular varieties, and of that peculiarly waved and corrugated variety simulating fossil wood, though really a mere form of laminated structure, which also occurs at Warrensburg, New York, and at the Marinski mine in Siberia. Many of the veins are not true fissures, but rather constitute a network of shrinkage cracks or segregation veins traversing in countless numbers the containing rock, and most irregular in their dimensions, so that they often resemble strings of nodular masses. It is most probable that the graphite of the veins was originally introduced as a liquid or plastic hydrocarbon; but in whatever way introduced, the character of the veins indicates that in the case of the greater number of them the carbonaceous material must have been derived from the bedded rocks traversed by these veins, to which it bears the same relation with the veins of bitumen found in the bituminous shales of the Carboniferous and Silurian rocks. Nor can there be any doubt that the graphite found in the beds has been deposited along with the calcareous matter or muddy and sandy sediment of which these beds were originally composed.
"Geology of Canada," 1863.
Paper by the author on Laurentian Graphite, "Journal of London Geological Society," 1876.
The quantity of graphite in the Lower Laurentian series is enormous. Some years ago, in the township of Buckingham, on the Ottawa River, I examined a band of limestone believed to be a continuation of that described by Sir W. E. Logan as the Green Lake limestone. It was estimated to amount, with some thin interstratified bands of gneiss, to a thickness of six hundred feet or more, and was found to be filled with disseminated crystals of graphite and veins of the mineral to such an extent as to constitute in some places one-fourth of the whole; and, making every allowance for the poorer portions, this band cannot contain in all a less vertical thickness of pure graphite than from twenty to thirty feet. In the adjoining township of Lochaber Sir W. E. Logan notices a band from twenty-five to thirty feet thick, reticulated with graphite veins to such an extent as to be mined with profit for the mineral. At another place in the same district a bed of graphite from ten to twelve feet thick, and yielding 20 per cent, of the pure material, is worked. As it appears in the excavation made by the quarrymen, it resembled a bed of coal; and a block from this bed, about four feet thick, was a prominent object in the Canadian department of the Colonial Exhibition of 1886. When it is considered that graphite occurs in similar abundance at several other horizons, in beds of limestone which have been ascertained by Sir W. E. Logan to have an aggregate thickness of thirty-five hundred feet, it is scarcely an exaggeration to maintain that the quantity of carbon in the Laurentian is equal to that in similar areas of the Carboniferous system. It is also to be observed that an immense area in Canada appears to be occupied by these graphitic and Eozoon limestones, and that rich graphitic deposits exist in the continuation of this system in the State of New York, while in rocks believed to be of this age near St. John, New Brunswick, there is a very thick bed of graphitic limestone, and associated with it three regular beds of graphite, having an aggregate thickness of about five feet.
Matthew in "Quarterly Journal of the Geological Society," vol. xxi., p. 423. "Acadian Geology," p. 662.
It may fairly be assumed that in the present world, and in those geological periods with whose organic remains we are more familiar than with those of the Laurentian, there is no other source of unoxidized carbon in rocks than that furnished by organic matter, and that this has obtained its carbon in all cases, in the first instance, from the deoxidation of carbonic acid by living plants. No other source of carbon can, I believe, be imagined in the Laurentian period. We may, however, suppose either that the graphitic matter of the Laurentian has been accumulated in beds like those of coal, or that it has consisted of diffused bituminous matter similar to that in more modern bituminous shales and bituminous and oil-bearing limestones. The beds of graphite near St. John, some of those in the gneiss at Ticonderoga in New York, and at Lochaber and Buckingham, and elsewhere in Canada, are so pure and regular that one might fairly compare them with the graphitic coal of Rhode Island. These instances, however, are exceptional, and the greater part of the disseminated and vein graphite might rather be likened in its mode of occurrence to the bituminous matter in bituminous shales and limestones.
We may compare the disseminated graphite to that which we find in those districts of Canada in which Silurian and Devonian bituminous shales and limestones have been metamorphosed and converted into graphitic rocks not very dissimilar to those in the less altered portions of the Laurentian. In like manner it seems probable that the numerous reticulating veins of graphite may have been formed by the segregation of bituminous matter into fissures and planes of least resistance, in the manner in which such veins occur in modern bituminous limestones and shales. Such bituminous veins occur in the Lower Carboniferous limestone and shale of Dorchester and Hillsborough, New Brunswick, with an arrangement very similar to that of the veins of graphite; and in the Quebec rocks of Point Levi, veins attaining to a thickness of more than a foot, are filled with a coaly matter having a transverse columnar structure, and regarded by Logan and Hunt as an altered bitumen. These palæozoic analogies would lead us to infer that the larger part of the Laurentian graphite falls under the second class of deposits above mentioned, and that, if of vegetable origin, the organic matter must have been thoroughly disintegrated and bituminised before it was changed into graphite. This would also give a probability that the vegetation implied was aquatic, or at least that it was accumulated under water.
Granby, Melbourne, Owl's Head, &c., "Geology of Canada," 1863, p. 599.
Dr. Hunt has, however, observed an indication of terrestrial vegetation, or at least of subaërial decay, in the great beds of Laurentian iron-ore. These, if formed in the same manner as more modern deposits of this kind, would imply the reducing and solvent action of substances produced in the decay of plants. In this case such great ore-beds as that of Hull, on the Ottawa, seventy feet thick, or that near Newborough, two hundred feet thick, must represent a corresponding quantity of vegetable matter which has totally disappeared. It may be added that similar demands on vegetable matter as a deoxidising agent are made by the beds and veins of metallic sulphides of the Laurentian, though some of the latter are no doubt of later date than the Laurentian rocks themselves.
"Geology of Canada," 1863.
It would be very desirable to confirm such conclusions as those above deduced by the evidence of actual microscopic structure. It is to be observed, however, that when, in more modern sediments, Algæ have been converted into bituminous matter, we cannot ordinarily obtain any structural evidence of the origin of such bitumen, and in the graphitic slates and limestones derived from the metamorphosis of such rocks no organic structure remains. It is true that, in certain bituminous shales and limestones of the Silurian system, shreds of organic tissue can sometimes be detected, and in some cases, as in the Lower Silurian limestone of the La Cloche Mountains in Canada, the pores of brachiopodous shells and the cells of corals have been penetrated by black bituminous matter, forming what may be regarded as natural injections, sometimes of much beauty. In correspondence with this, while in some Laurentian graphitic rocks, as, for instance, in the compact graphite of Clarendon, the carbon presents a curdled appearance due to segregation, and precisely similar to that of the bitumen in more modern bituminous rocks, I can detect in the graphitic limestones occasional fibrous structures which may be remains of plants, and in some specimens vermicular lines, which I believe to be tubes of Eozoon penetrated by matter once bituminous, but now in the state of graphite.
When palæozoic land-plants have been converted into graphite, they sometimes perfectly retain their structure. Mineral charcoal, with structure, exists in the graphitic coal of Rhode Island. The fronds of ferns, with their minutest veins perfect, are preserved in the Devonian shales of St. John, in the state of graphite; and in the same formation there are trunks of Conifers (Dadoxylon Ouangondianum) in which the material of the cell-walls has been converted into graphite, while their cavities have been filled with calcareous spar and quartz, the finest structures being preserved quite as well as in comparatively unaltered specimens from the coal-formation. No structures so perfect have as yet been detected in the Laurentian, though in the largest of the three graphitic beds at St. John there appear to be fibrous structures, which I believe may indicate the existence of land-plants. This graphite is composed of contorted and slickensided laminæ, much like those of some bituminous shales and coarse coals; and in these are occasional small pyritous masses which show hollow carbonaceous fibres, in some cases presenting obscure indications of lateral pores. I regard these indications, however, as uncertain; and it is not as yet fully ascertained that these beds at St. John are on the same geological horizon with the Lower Laurentian of Canada, though they certainly underlie the Primordial series of the Acadian group, and are separated from it by beds having the character of the Huronian.
"Acadian Geology," p. 535. In calcined specimens the structures remain in the graphite after decalcification by an acid.
There is thus no absolute impossibility that distinct organic tissues may be found in the Laurentian graphite, if formed from land-plants, more especially if any plants existed at that time having true woody or vascular tissues; but it cannot with certainty be affirmed that such tissues have been found. It is possible, however, that in the Laurentian period the vegetation of the land may have consisted wholly of cellular plants, as, for example, mosses and lichens; and if so, there would be comparatively little hope of the distinct preservation of their forms or tissues, or of our being able to distinguish the remains of land-plants from those of Algæ.
We may sum up these facts and considerations in the following statements: First, that somewhat obscure traces of organic structure can be detected in the Laurentian graphite; secondly, that the general arrangement and microscopic structure of the substance corresponds with that of the carbonaceous and bituminous matters in marine formations of more modern date; thirdly, that if the Laurentian graphite has been derived from vegetable matter, it has only undergone a metamorphosis similar in kind to that which organic matter in metamorphosed sediments of later age has experienced; fourthly, that the association of the graphitic matter with organic limestone, beds of iron-ore, and metallic sulphides greatly strengthens the probability of its vegetable origin; fifthly, that when we consider the immense thickness and extent of the Eozoonal and graphitic limestones and iron-ore deposits of the Laurentian, if we admit the organic origin of the limestone and graphite, we must be prepared to believe that the life of that early period, though it may have existed under low forms, was most copiously developed, and that it equalled, perhaps surpassed, in its results, in the way of geological accumulation, that of any subsequent period.
Many years ago, at the meeting of the American Association in Albany, the writer was carrying into the room of the Geological Section a mass of fossil wood from the Devonian of Gaspé, when he met the late Professor Agassiz, and remarked that the specimen was the remains of a Devonian tree contemporaneous with his fishes of that age. "How I wish I could sit under its shade!" was the smiling reply of the great zoölogist; and when we think of the great accumulations of Laurentian carbon, and that we are entirely ignorant of the forms and structures of the vegetation which produced it, we can scarcely suppress a feeling of disappointment. Some things, however, we can safely infer from the facts that are known, and these it may be well to mention.
The climate and atmosphere of the Laurentian may have been well adapted for the sustenance of vegetable life. We can scarcely doubt that the internal heat of the earth still warmed the waters of the sea, and these warm waters must have diffused great quantities of mists and vapours over the land, giving a moist and equable if not a very clear atmosphere. The vast quantities of carbon dioxide afterwards sealed up in limestones and carbonaceous beds must also have still floated in the atmosphere and must have supplied abundance of the carbon, which constitutes the largest ingredient in vegetable tissues. Under these circumstances the whole world must have resembled a damp, warm greenhouse, and plants loving such an atmosphere could have grown luxuriantly. In these circumstances the lower forms of aquatic vegetation and those that love damp, warm air and wet soil would have been at home.
If we ask more particularly what kinds of plants might be expected to be introduced in such circumstances, we may obtain some information from the vegetation of the succeeding Palæozoic age, when such conditions still continued to a modified extent. In this period the club-mosses, ferns, and mare's-tails engrossed the world and grew to sizes and attained degrees of complexity of structure not known in modern times. In the previous Laurentian age something similar may have happened to Algæ, to Fungi, to Lichens, to Liverworts, and Mosses. The Algæ may have attained to gigantic dimensions, and may have even ascended out of the water in some of their forms. These comparatively simple cellular and tubular structures, now degraded to the humble position of flat lichens or soft or corky fungi, or slender cellular mosses, may have been so strengthened and modified as to constitute forest-trees. This would be quite in harmony with what is observed in the development of other plants in primitive geological times; and a little later in this history we shall see that there is evidence in the flora of the Silurian of a survival of such forms.
It may be that no geologist or botanist will ever be able to realise these dreams of the past. But, on the other hand, it is quite possible that some fortunate chance may have somewhere preserved specimens of Laurentian plants showing their structure.
In any case we have here presented to us the strange and startling fact that the remarkable arrangement of protoplasmic matter and chlorophyll, which enables the vegetable cell to perform, with the aid of solar light, the miracle of decomposing carbon dioxide and water, and forming with them woody and corky tissues, had already been introduced upon the earth. It has been well said that no amount of study of inorganic nature would ever have enabled any one to anticipate the possibility of the construction of an apparatus having the chemical powers of the living vegetable cell. Yet this most marvellous structure seems to have been introduced in the full plenitude of its powers in the Laurentian age.
Whether this early Laurentian vegetation was the means of sustaining any animal life other than marine Protozoa, we do not know. It may have existed for its own sake alone, or merely as a purifier of the atmosphere, in preparation for the future introduction of land-animals. The fact that there have existed, even in modern times, oceanic islands rich in vegetation, yet untenanted by the higher forms of animal life, prepares us to believe that such conditions may have been general or universal in the primeval times we are here considering.
If we ask to what extent the carbon extracted from the atmosphere and stored up in the earth has been, or is likely to be, useful to man, the answer must be that it is not in a state to enable it to be used as mineral fuel. It has, however, important uses in the arts, though at present the supply seems rather in excess of the demand, and it may well be that there are uses of graphite still undiscovered, and to which it will yet be applied.
Finally, it is deserving of notice that, if Laurentian graphite indicates vegetable life, it indicates this in vast profusion. That incalculable quantities of vegetable matter have been oxidised and have disappeared we may believe on the evidence of the vast beds of iron-ore; and, in regard to that preserved as graphite, it is certain that every inch of that mineral must indicate many feet of crude vegetable matter.
It is remarkable that, in ascending from the Laurentian, we do not at first appear to advance in evidences of plant-life. The Huronian age, which succeeded the Laurentian, seems to have been a disturbed and unquiet time, and, except in certain bands of iron-ore and some dark slates coloured with carbonaceous matter, we find in it no evidence of vegetation. In the Cambrian a great subsidence of our continents began, which went on, though with local intermissions and reversals, all through the Siluro-Cambrian or Ordovician time. These times were, for this reason, remarkable for the great abundance and increase of marine animals rather than of land-plants. Still, there are some traces of land vegetation, and we may sketch first the facts of this kind which are known, and then advert to some points relating to the earlier Algæ, or sea-weeds.
An eminent Swedish geologist, Linnarsson, has described, under the name of Eophyton, certain impressions on old Cambrian rocks in Sweden, and which certainly present very plant-like forms. They want, however, any trace of carbonaceous matter, and seem rather to be grooves or marks cut in clay by the limbs or tails of some aquatic animal, and afterwards filled up and preserved by succeeding deposits. After examining large series of these specimens from Sweden, and from rocks of similar age in Canada, I confess that I have no faith in their vegetable nature.
The oldest plants known to me, and likely to have been of higher grade than Algæ, are specimens kindly presented to me by Dr. Alleyne Nicholson, of Aberdeen, and which he had named Buthotrephis Harknessii and B. radiata. They are from the Skiddaw rocks of Cumberland. On examining these specimens, and others subsequently collected in the same locality by Dr. Gr. M. Dawson, while convinced by their form and carbonaceous character that they are really plants, I am inclined to refer them not to Algæ, but probably to Rhizocarps. They consist of slender branching stems, with whorls of elongate and pointed leaves, resembling the genus Annularia of the coal formation. I am inclined to believe that both of Nicholson's species are parts of one plant, and for this I have proposed the generic name Protannularia (Fig. 1). Somewhat higher in the Siluro-Cambrian, in the Cincinnati group of America, Lesquereux has found some minute radiated leaves, referred by him to the genus Sphenophyllum, which is also allied to Rhizocarps. Still more remarkable is the discovery in the same beds of a stem with rhombic areoles or leaf-bases, to which the name Protostigma has been given. If a plant, this may have been allied to the club-mosses. This seems to be all that we at present know of land-vegetation in the Siluro-Cambrian. So far as the remains go, they indicate the presence of the families of Rhizocarps and of Lycopods.
"Geological Magazine," 1869.
See figure in next chapter.
Protostigma sigillarioides, Lesquereux.
If we ascend into the Upper Silurian, or Silurian proper, the evidences of land vegetation somewhat increase. In 1859 I described, in "The Journal of the Geological Society" of London, a remarkable tree from the Lower Erian of Gaspé, under the name Prototaxites, but for which I now prefer the name Nematophyton. When in London, in 1870, I obtained permission to examine certain specimens of spore-cases or seeds from the Upper Ludlow (Silurian) formation of England, and which had been described by Sir Joseph Hooker under the name Pachytheca. In the same slabs with these I found fragments of fossil wood identical with those of the Gaspé plant. Still later I recognised similar fragments associated also with Pachytheca in the Silurian of Cape Bon Ami, New Brunswick. Lastly, Dr. Hicks has discovered similar wood, and also similar fruits, in the Denbighshire grits, at the base of the Silurian.
"Journal of the Geological Society," August, 1881.
Figs. 2, 3, and 4 are drawn from nature by Prof. Penhallow, of McGill College.
From comparison of this singular wood, the structure of which is represented in Figs. 2, 3, 4, with the débris of fossil taxine woods, mineralised after long maceration in water, I was inclined to regard Prototaxites, or, as I have more recently named it, Nematophyton, as a primeval gymnosperm allied to those trees which Unger had described from the Erian of Thuringia, under the name Aporoxylon. Later examples of more lax tissues from branches or young stems, and the elaborate examinations kindly undertaken for me by Professor Penhallow and referred to in a note to this chapter, have induced me to modify this view, and to hold that the tissues of these singular trees, which seem to have existed from the beginning of the Silurian age and to have finally disappeared in the early Erian, are altogether distinct from any form of vegetation hitherto known, and are possibly survivors of that prototypal flora to which I have already referred. They are trees of large size, with a coaly bark and large spreading roots, having the surface of the stem smooth or irregularly ribbed, but with a nodose or jointed appearance. Internally, they show a tissue of long, cylindrical tubes, traversed by a complex network of horizontal tubes thinner walled and of smaller size. The tubes are arranged in concentric zones, which, if annual rings, would in some specimens indicate an age of one hundred and fifty years. There are also radiating spaces, which I was at first disposed to regard as true medullary rays, or which at least indicate a radiating arrangement of the tissue. They now seem to be spaces extending from the centre towards the circumference of the stem, and to have contained bundles of tubes gathered from the general tissue and extending outward perhaps to organs or appendages on the surface. Carruthers has suggested a resemblance to Algæ, and has even proposed to change the name to Nematophycus, or "thread-sea-weed"; but the resemblance is by no means clear, and it would be quite as reasonable to compare the tissue to that of some Fungi or Lichens, or even to suppose that a plant composed of cylindrical tubes has been penetrated by the mycelium or spawn of a dry-rot fungus. But the tissues are too constant and too manifestly connected with each other to justify this last supposition. That the plant grew on land I cannot doubt, from its mode of occurrence; that it was of durable and resisting character is shown by its state of preservation; and the structure of the seeds called Pachytheca, with their constant association with these trees, give countenance to the belief that they are the fruit of Nematophyton. Of the foliage or fronds of these strange plants we unfortunately know nothing. They seem, however, to realise the idea of arboreal plants having structures akin to those of thallophytes, but with seeds so large and complex that they can scarcely be regarded as mere spores. They should perhaps constitute a separate class or order to which the name Nematodendreæ may be given, and of which Nematophyton will constitute one genus and Aporoxylon of Unger another.
"Palæontologie des Thuringer Waldes," 1856.
See report by the author on "Erian Flora of Canada," 1871 and 1882, for full description of these fossils.
Another question arises as to the possible relation of these plants to other trees known by their external forms. The Protostigma of Lesquereux has already been referred to, and Claypole has described a tree from the Clinton group of the United States, with large ovate leaf-bases, to which he has given the name Glyptodendron. If the markings on these plants are really leaf-bases, they can scarcely have been connected with Nematophyton, because that tree shows no such surface-markings, though, as we have seen, it had bundles of tubes passing diagonally to the surface. These plants were more probably trees with an axis of barred vessels and thick, cellular bark, like the Lepidodendron of later periods, to be noticed in the sequel. Dr. Hicks has also described from the same series of beds which afforded the fragments of Nematophyton certain carbonised dichotomous stems, which he has named Berwynia. It is just possible that these plants may have belonged to the Nematodendreæ. The thick and dense coaly matter which they show resembles the bark of these trees, the longitudinal striation in some of them may represent the fibrous structure, and the lateral projections which have been compared to leaves or leaf-bases may correspond with the superficial eminences of Nematophyton, and the spirally arranged punctures which it shows on its surface. In this case I should be disposed to regard the supposed stigmaria-like roots as really stems, and the supposed rootlets as short, spine-like rudimentary leaves. All such comparisons must, however, in the mean time be regarded as conjectural. We seem, however, to have here a type of tree very dissimilar to any even of the later Palæozoic age, which existed throughout the Silurian, and probably further back, which ceased to exist early in the Erian age, and before the appearance of the ordinary coniferous and lepidodendroid trees. May it not have been a survivor of an old arboreal flora extending back even to the Laurentian itself?
"American Journal of Science," 1878.
Multitudes of markings occurring on the surfaces of the older rocks have been referred to the Algæ or sea-weeds, and indeed this group has been a sort of refuge for the destitute to which palæontologists have been accustomed to refer any anomalous or inexplicable form which, while probably organic, could not be definitely referred to the animal kingdom. There can be no question that some of these are truly marine plants; and that plants of this kind occur in formations older than those in which we first find land-plants, and that they have continued to inhabit the sea down to the present time. It is also true that the oldest of these Algæ closely resemble in form plants of this kind still existing; and, since their simple cellular structures and soft tissues are scarcely ever preserved, their general forms are all that we can know, so that their exact resemblance to or difference from modern types can rarely be determined. For the same reasons it has proved difficult clearly to distinguish them from mere inorganic markings or the traces of animals, and the greatest divergence of opinion has occurred in recent times on these subjects, as any one can readily understand who consults the voluminous and well-illustrated memoirs of Nathorst, Williamson, Saporta, and Delgado.
The author of this work has given much attention to these remains, and has not been disposed to claim for the vegetable kingdom so many of them as some of his contemporaries. The considerations which seem most important in making such distinctions are the following: 1. The presence or absence of carbonaceous matter. True Algæ not infrequently present at least a thin film of carbon representing their organic matter, and this is the more likely to occur in their case, as organic matters buried in marine deposits and not exposed to atmospheric oxidation are very likely to be preserved. 2. In the absence of organic matter, the staining of the containing rock, the disappearance or deoxidation of its ferruginous colouring matter, or the presence of iron pyrite may indicate the removal of organic matter by decay. 3. When organic matter and indications of it are altogether absent, and form alone remains, we have to distinguish from Algæ, trails and burrows similar to those of aquatic animals, casts of shrinkage-cracks, water-marks, and rill-marks widely diffused over the surfaces of beds. 4. Markings depressed on the upper surfaces of beds, and filled with the material of the succeeding layer, are usually mere impressions. The cases of possible exceptions to this are very rare. On the contrary, there are not infrequently forms in relief on the surfaces of rocks which are not Algæ, but may be shallow burrows arched upward on top, or castings of worms thrown up upon the surface. Sometimes, however, they may have been left by denudation of the surrounding material, just as footprints on dry snow remain in relief after the surrounding loose material has been drifted away by the wind; the portion consolidated by pressure being better able to resist the denuding agency.
"Impressions and Footprints of Aquatic Animals," "American Journal of Science," 1873.
The footprints from the Potsdam sandstone in Canada, for which the name Protichnites was proposed by Owen, and which were by him referred to crustaceans probably resembling Limulus, were shown by the writer, in 1862, to correspond precisely with those of the American Limulus (Polyphemus Occidentalis) (Fig. 5). I proved by experiment with the modern animal that the recurring series of groups of markings were produced by the toes of the large posterior thoracic feet, the irregular scratches seen in Protichnites lineatus by the ordinary feet, and the central furrow by the tail. It was also shown that when the Limulus uses its swimming-feet it produces impressions of the character of those named Climactichnites, from the same beds which afford Protichnites. The principal difference between Protichnites and their modern representatives is that the latter have two lateral furrows produced by the sides of the carapace, which are wanting in the former.
"Canadian Naturalist," vol. vii.
I subsequently applied the same explanation to several other ancient forms now known under the general name Bilobites (Figs. 6 and 7).
The name Bilobites was originally proposed by De Kay for a bivalve shell (Conocardium). Its application to supposed Algæ was an error, but this is of the less consequence, as these are not true plants but only animal trails.
The tuberculated impressions known as Phymatoderma and Caulerpites may, as Zeiller has shown, be made by the burrowing of the mole-cricket, and fine examples occurring in the Clinton formation of Canada are probably the work of Crustacea. It is probable, however, that some of the later forms referred to these genera are really Algæ related to Caulerpa, or even branches of Conifers of the genus Brachyphyllum.
Nereites and Planulites are tracks and burrows of worms, with or without marks of setæ, and some of the markings referred to Palæochorda, Palæophycus, and Scolithus have their places here. Many examples highly illustrative of the manner of formation of the impressions are afforded by Canadian rocks (Fig. 8).
Branching forms referred to Licrophycus of Billings, and some of those referred to Buthotrephis, Hall, as well as radiating markings referable to Scotolithus, Gyrophyllites, and Asterophycus, are explained by the branching burrows of worms illustrated by Nathorst and the author. Astropolithon, a singular radiating marking of the Canadian Cambrian, seems to be something organic, but of what nature is uncertain (Fig. 9).
Supplement to "Acadian Geology."
Rhabdichnites and Eophyton belong to impressions explicable by the trails of drifting sea-weeds, the tail-markings of Crustacea, and the ruts ploughed by bivalve mollusks, and occurring in the Silurian, Erian, and Carboniferous rocks. Among these are the singular bilobate forms described as Rusophycus by Hall, and which are probably burrows or resting-places of crustaceans. The tracks of such animals, when walking, are the jointed impressions known as Arthrophycus and Crusiana. I have shown by the mode of occurrence of these, and Nathorst has confirmed this conclusion by elaborate experiments on living animals, that these forms are really trails impressed on soft sediments by animals and mostly by crustaceans.
"Canadian Naturalist," 1864.
I agree with Dr. Williamson in believing that all or nearly all the forms referred to Crossochorda of Schimper are really animal impressions allied to Nereites, and due either to worms or, as Nathorst has shown to be possible, to small crustaceans. Many impressions of this kind occur in the Silurian beds of the Clinton series in Canada and New York, and are undoubtedly mere markings.
"Tracks from Yoredale Rocks," "Manchester Literary and Philosophical Society," 1885.
It is worthy of note that these markings strikingly resemble the so-called Eophyton, described by Torell from the Primordial of Sweden, and by Billings from that of Newfoundland; and which also occur abundantly in the Primordial of New Brunswick. After examining a series of these markings from Sweden shown to me by Mr. Carruthers in London, and also specimens from Newfoundland and a large number in situ at St. John, I am convinced that they cannot be plants, but must be markings of the nature of Rhabdichnites. This conclusion is based on the absence of carbonaceous matter, the intimate union of the markings with the surface of the stone, their indefinite forms, their want of nodes or appendages, and their markings being always of such a nature as could be produced by scratches of a sharp instrument. Since, however, fishes are yet unknown in beds of this age, they may possibly be referred to the feet or spinous tails of swimming crustaceans. Salter has already suggested this origin for some scratches of somewhat different form found in the Primordial of Great Britain. He supposed them to have been the work of species of Hymenocaris. These marks may, however, indicate the existence of some free-swimming animals of the Primordial seas as yet unknown to us.
Three other suggestions merit consideration in this connection. One is that Algæ and also land-plants, drifting with tides or currents, often make the most remarkable and fantastic trails. A marking of this kind has been observed by Dr. G. M. Dawson to be produced by a drifted Laminaria, and in complexity it resembled the extraordinary Ænigmichnus multiformis of Hitchcock from the Connecticut sandstones. Much more simple markings of this kind would suffice to give species of Eophyton. Another is furnished by a fact stated to the author by Prof. Morse, namely, that Lingulæ, when dislodged from their burrows, trail themselves over the bottom like worms, by means of their cirri. Colonies of these creatures, so abundant in the Primordial, may, when obliged to remove, have covered the surfaces of beds of mud with vermicular markings. The third is that the Rhabdichnite-markings resemble some of the grooves in Silurian rocks which have been referred to trails of Gasteropods, as, for instance, those from the Clinton group, described by Hall.
Another kind of markings not even organic, but altogether depending on physical causes, are the beautiful branching rill-marks produced by the oozing of water out of mud and sand-banks left by the tide, and which sometimes cover great surfaces with the most elaborate tracery, on the modern tidal shores as well as in some of the most ancient rocks. Dendrophycus of Lesquereux seems to be an example of rill-mark, as well as Aristophycus, Cloephycus, and Zygopliycus, of Miller and Dyer, from the Lower Silurian.
"Coal Flora of Pennsylvania," vol. iii., Plate 88.
Rill-marks occur in very old rocks, but are perhaps most beautifully preserved in the Carboniferous shales and argillaceous sandstones, and even more elaborately on the modern mud-banks of the Bay of Fundy. Some of these simulate ferns and fronds of Laminariæ, and others resemble roots, fucoids allied to Buthotrephis, or the radiating worm-burrows already referred to (Fig. 10).
"Journal of the Geological Society," vol. xii., p. 251.
"Acadian Geology," 2d ed., p. 26.
Shrinkage-cracks are also abundant in some of the Carboniferous beds, and are sometimes accompanied with impressions of rain-drops. When finely reticulated they might be mistaken for the venation of leaves, and, when complicated with little rill-marks tributary to their sides, they precisely resemble the Dictyolites of Hall from the Medina sandstone (Fig. 11).
An entirely different kind of shrinkage-crack is that which occurs in certain carbonised and flattened plants, and which sometimes communicates to them a marvellous resemblance to the netted under surface of an exogenous leaf. Flattened stems of plants and layers of cortical matter, when carbonised, shrink in such a manner as to produce minute reticulated cracks. These become filled with mineral matter before the coaly substance has been completely consolidated. A further compression occurs, causing the coaly substance to collapse, leaving the little veins of harder mineral matter projecting. These impress their form upon the clay or shale above and below, and thus when the mass is broken open we have a carbonaceous film or thin layer covered with a network of raised lines, and corresponding minute depressed lines on the shale in contact with it. The reticulations are generally irregular, but sometimes they very closely resemble the veins of a reticulately veined leaf. One of the most curious specimens in my possession was collected by Mr. Elder in the Lower Carboniferous of Horton Bluff. The little veins which form the projecting network are in this case white calcite; but at the surface their projecting edges are blackened with a carbonaceous film.
Slickensided bodies, resembling the fossil fruits described by Geinitz as Gulielmites, and the objects believed by Fleming and Carruthers to be casts of cavities filled with fluid, abound in the shales of the Carboniferous and Devonian. They are, no doubt, in most cases the results of the pressure and consolidation of the clay around small solid bodies, whether organic, fragmentary, or concretionary. They are, in short, local slickensides precisely similar to those found so plentifully in the coal under-clays, and which, as I have elsewhere shown, resulted from the internal giving way and slipping of the mass as the roots of Stigmaria decayed within it. Most collectors of fossil plants in the older formations must, I presume, be familiar with appearances of this kind in connection with small stems, petioles, fragments of wood, and carpolites. I have in my collection petioles of ferns and fruits of the genus Trigonocarpum partially slickensided in this way, and which if wholly covered by this kind of marking could scarcely have been recognised. I have figured bodies of this kind in my report on the Devonian and Upper Silurian plants of Canada, believing them, owing to their carbonaceous covering, to be probably slickensided fruits, though of uncertain nature. In every case I think these bodies must have had a solid nucleus of some sort, as the severe pressure implied in slickensiding is quite incompatible with a mere "fluid-cavity," even supposing this to have existed.
"Journal of the Geological Society," June, 1871.
Ibid., vol. x., p. 14.
Prof. Marsh has well explained another phase of the influence of hard bodies in producing partial slickensides, in his paper on Stylolites, read before the American Association in 1867, and the application of the combined forces of concretionary action and slickensiding to the production of the cone-in-cone concretions, which occur in the coal-formation and as low as the Primordial. I have figured a very perfect and beautiful form of this kind from the coal-formation of Nova Scotia, which is described in "Acadian Geology" (Fig. 12).
I have referred to these facts here because they are relatively more important in that older period, which may be named the age of Algæ, and because their settlement now will enable us to dispense with discussions of this kind further on. The able memoirs of Nathorst and Williamson should be studied by those who desire further information.
Appendix, p. 676, edition of 1878.
But it may be asked, "Are there no real examples of fossil Algæ?" I believe there are many such, but the difficulty is to distinguish them. Confining ourselves to the older rocks, the following may be noted:
The genus Buthotrephis of Hall, which is characterised as having stems, sub-cylindric or compressed, with numerous branches, which are divaricating and sometimes leaf-like, contains some true Algæ. Hall's B. gracilis, from the Siluro-Cambrian, is one of these. Similar plants, referred to the same species, occur in the Clinton and Niagara formations, and a beautiful species, collected by Col. Grant, of Hamilton, and now in the McGill College collection, represents a broader and more frondose type of distinctly carbonaceous character. It may be described as follows:
Buthotrephis Grantii, S. N. (Fig. 13).--Stems and fronds smooth and slightly striate longitudinally, with curved and interrupted striæ. Stem thick, bifurcating, the divisions terminating in irregularly pinnate fronds, apparently truncate at the extremities. The quantity of carbonaceous matter present would indicate thick, though perhaps flattened, stems and dense fleshy fronds.
The species Buthotrephis subnodosa and B. flexuosa, from the Utica shale, are also certainly plants, though it is possible, if their structures and fruit were known, some of these might be referred to different genera. All of these plants have either carbonaceous matter or produce organic stains on the matrix.
The organism with diverging wedge-shaped fronds, described by Hall as Sphenothallus angustifolius, is also a plant. Fine specimens, in the collection of the Geological Survey of Canada, show distinct evidence of the organic character of the wedge-shaped fronds. It is from the Utica shale, and elsewhere in the Siluro-Cambrian. It is just possible, as suggested by Hall, that this plant may be of higher rank than the Algæ.
The genus Palæophycus of Hall includes a great variety of uncertain objects, of which only a few are probably true Algæ. I have specimens of fragments similar to his P. virgatus, which show distinct carbonaceous films, and others from the Quebec group, which seem to be cylindrical tubes now flattened, and which have contained spindle-shaped sporangia of large size. Tortuous and curved flattened stems, or fronds, from the Upper Silurian limestone of Gaspé, also show organic matter.
Respecting the forms referred to Licrophycus by Billings, containing stems or semi-cylindrical markings springing from a common base, I have been in great doubt. I have not seen any specimens containing unequivocal organic matter, and am inclined to think that most of them, if not the whole, are casts of worm-burrows, with trails radiating from them.
Though I have confined myself in this notice to plants, or supposed plants, of the Lower Palæozoic, it may be well to mention the remarkable Cauda-Galli fucoids, referred by Hall to the genus Spirophyton, and which are characteristic of the oldest Erian beds. The specimens which I have seen from New York, from Gaspé, and from Brazil, leave no doubt in my mind that these were really marine plants, and that the form of a spiral frond, assigned to them by Hall, is perfectly correct. They must have been very abundant and very graceful plants of the early Erian, immediately after the close of the Silurian period.
We come now to notice certain organisms referred to Algæ, and which are either of animal origin, or are of higher grade than the sea-weeds. We have already discussed the questions relating to Prototaxites. Drepanophycus, of Goeppert, I suspect, is only a badly preserved branch or stem of the Erian land-plant known as Arthrostigma. In like manner, Haliserites Dechenianus, of Goeppert, is evidently the land-plant known as Psilophyton. Sphærococcites dentatus and S. serra--the Fucoides dentatus and serra of Brongniart, from Quebec--are graptolites of two species quite common there. Dictyophyton and Uphantenia, as described by Hall and the author, are now known to be sponges. They have become Dictyospongiæ. The curious and very ancient; fossils referred by Forbes to the genus Oldhamia are perhaps still subject to doubt, but are usually regarded as Zoöphytes, though it is quite possible they may be plants. Though I have not seen the specimens, I have no doubt whatever that the plants, or the greater part of them, from the Silurian of Bohemia, described by Stur as Algæ and Characeæ, are really land-plants, some of them of the genus Psilophyton. I may say in this connection that specimens of flattened Psilophyton and Arthrostigma, in the Upper Silurian and Erian of Gaspé, would probably have been referred to Algæ, but for the fact that in some of them the axis of barred vessels is preserved.
"Fossile Flora," 1852, p. 92, Table xli.
Ibid., p. 88, Table ii.
Brongniart, "Vegeteaux Fossiles," Plate vi., Figs. 7 to 12.
"Proceedings of the Vienna Academy," 1881. Hostinella, of this author, is almost certainly Psilophyton, and his Barrandiana seems to include Arthrostigma, and perhaps leafy branches of Berwynia. These curious plants should be re-examined.
It is not surprising that great difficulties have occurred in the determination of fossil Algæ. Enough, however, remains certain to prove that the old Cambrian and Silurian seas were tenanted with sea-weeds not very dissimilar from those of the present time. It is further probable that some of the graphitic, carbonaceous, and bituminous shales and limestones of the Silurian owe their carbonaceous matters to the decomposition of Algæ, though possibly some of it may have been derived from Graptolites and other corneous Zoöphytes. In any case, such microscopic examinations of these shales as I have made, have not produced any evidence of the existence of plants of higher grade, while those of the Erian and Carboniferous periods, similar to the naked eye, abound in such evidence. It is also to be observed that, on the surfaces of beds of sandstone in the Upper Cambrian, carbonaceous débris, which seems to be the remains of either aquatic or land plants, is locally not infrequent.
Referring to the land vegetation of the older rocks, it is difficult to picture its nature and appearance. We may imagine the shallow waters filled with aquatic or amphibious Rhizocarpean plants, vast meadows or brakes of the delicate Psilophyton and the starry Protannularia and some tall trees, perhaps looking like gigantic club-mosses, or possibly with broad, flabby leaves, mostly cellular in texture, and resembling Algæ transferred to the air. Imagination can, however, scarcely realise this strange and grotesque vegetation, which, though possibly copious and luxuriant, must have been simple and monotonous in aspect, and, though it must have produced spores and seeds and even fruits, these were probably all of the types seen in the modern acrogens and gymnosperms.
"In garments green, indistinct in the twilight, They stand like Druids of old, with voices sad and prophetic."
Prophetic they truly were, as we shall find, of the more varied forests of succeeding times, and they may also help us to realise the aspect of that still older vegetation, which is fossilised in the Laurentian graphite; though it is not impossible that this last may have been of higher and more varied types, and that the Cambrian and Silurian may have been times of depression in the vegetable world, as they certainly were in the submergence of much of the land.
These primeval woods served at least to clothe the nakedness of the new-born land, and they may have sheltered and nourished forms of land-life still unknown to us, as we find as yet only a few insects and scorpions in the Silurian. They possibly also served to abstract from the atmosphere some portion of its superabundant carbonic acid harmful to animal life, and they stored up supplies of graphite, of petroleum, and of illuminating gas, useful to man at the present day. We may write of them and draw their forms with, the carbon which they themselves supplied.
* * * * *
NOTE TO CHAPTER II.
Examination of Prototaxites (Nematophyton), by Prof. Penhallow, of McGill University.
Prof. Penhallow, having kindly consented to re-examine my specimens, has furnished me with elaborate notes of his facts and conclusions, of which the following is a summary, but which it is hoped will be published in full:
"1. Concentric Layers.--The inner face of each of these is composed of relatively large tubes, having diameters from 13·6 to 34·6 micro-millimetres. The outer face has tubes ranging from 13·8 to 27·6 mm. The average diameter in the lower surface approaches to 34, that in the outer to 13·8. There is, however, no abrupt termination to the surface of the layers, though in some specimens they separate easily, with shining surfaces.
"2. Minute Structure.--In longitudinal sections the principal part of the structure consists of longitudinal tubes of indeterminate length, and round in cross-section. They are approximately parallel, but in some cases may be seen to bend sinuously, and are not in direct contact. Finer myceloid tubes, 5·33 mm. in diameter, traverse the structure in all directions, and are believed to branch off from the larger tubes. In a small specimen supposed to be a branch or small stem, and in which the vertical tubes are somewhat distant from one another, this horizontal system is very largely developed; but is less manifest in the older stems. The tubes themselves show no structure. The ray-like openings in the substance of the tissue are evidently original parts of the structure, but not of the nature of medullary rays. They are radiating spaces running outward in an interrupted manner or so tortuously that they appear to be interrupted in their course from the centre towards the surface. They show tubes turning into them, branching into them, and approximately horizontal, but tortuous. On the external surface of some specimens these radial spaces are represented by minute pits irregularly or spirally arranged. The transverse swellings of the stem show no difference of structure, except that the tubes or cells may be a little more tortuous, and a transverse film of coaly matter extends from the outer coaly envelope inwardly. This may perhaps be caused by some accident of preservation. The outer coaly layer shows tubes similar to those of the stem. The horizontal or oblique flexures of the large tubes seem to be mainly in the vicinity of the radial openings, and it is in entering these that they have been seen to branch."
It is possible that these tubes may be merely part of the stem attached to the bark, which seems to me to indicate the same dense cellular structure seen in the bark of Lepidodendra, etc.
The conclusions arrived at by Prof. Penhallow are as follows:
"1. The plant was not truly exogenous, and the appearance of rings is independent of the causes which determine the layers of growth in exogenous plants.
"2. The plant was possessed of no true bark. Whatever cortical layer was present was in all probability a modification of the general structure,
On these points I would reserve the considerations: 1. That there must have been some relation between the mode of growth of these great stems and their concentric rings; and, 2. That the evidence of a bark is as strong as in the case of any Palæozoic tree in which the bark is, as usual, carbonised.
"3. An intimate relation exists between the large tubular cells and the myceloid filaments, the latter being a system of small branches from the former; the branching being determined chiefly in certain special openings which simulate medullary rays.
"4. The specimens examined exhibit no evidence of special decay, and the structure throughout is of a normal character.
"5. The primary structure consists of large tubular cells without apparent terminations, and devoid of structural markings, with which is associated a secondary structure of myceloid filaments arising from the former.
"6. The structure of Nematophyton as a whole is unique; at least there is no plant of modern type with which it is comparable. Nevertheless, the loose character of the entire structure; the interminable cells; their interlacing; and, finally, their branching into a secondary series of smaller filaments, point with considerable force to the true relationship of the stem as being with Algæ or other Thallophytes rather than with Grymnosperms. A more recent examination of a laminated resinous substance found associated with the plant shows that it is wholly amorphous, and, as indicated by distinct lines of flow, that it must have been in a plastic state at a former period. The only evidence of structure was found in certain well-defined mycelia, which may have been derived from associated vegetable matter upon which they were growing, and over which the plastic matrix flowed."
I have only to add to this description that when we consider that Nematophyton Logani was a large tree, sometimes attaining a diameter of more than two feet, and a stature of at least twenty before branching; that it had great roots, and gave off large branches; that it was an aërial plant, probably flourishing in the same swampy flats with Psilophyton, Arthrostigma, and Leptophleum; that the peculiar bodies known as Pachytheca were not unlikely its fruit--we have evidence that there were, in the early Palæozoic period, plants scarcely dreamt of by modern botany. Only when the appendages of these plants are more fully known can we hope to understand them. In the mean time, I may state that there were probably different species of these trees, indicated more particularly by the stems I have described as Nematoxylon and Celluloxylon There were, I think, some indications that the plants described by Carruthers as Berwynia, may also be found to have been generically the same. The resinous matter mentioned by Prof. Penhallow is found in great abundance in the beds containing Nematophyton, and must, I think, have been an exudation from its bark.
"Journal Geol. Society of London," 1863, 1881.
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