MARKINGS, FOOTPRINTS AND FUCOIDS.

[138]The discovery of the shells in this tree was made by Albert I. Hill, C.E. The tree is in Group XXVI. of Division 4 of my Joggins section. The original reptiliferous trees are in Group XV., and the lowest bed in Group VIII.

[138]The discovery of the shells in this tree was made by Albert I. Hill, C.E. The tree is in Group XXVI. of Division 4 of my Joggins section. The original reptiliferous trees are in Group XV., and the lowest bed in Group VIII.

If we exclude the allegedPalæorbisreferred to below, all the Palæozoic Pulmonifera hitherto found are American. Since, however, in the Carboniferous age, Batrachians, Arachnidans, Insects and Millipedes occur on both continents, it is not unlikely that ere long European species of land snails will be announced The species hitherto found in Eastern America are in every way strangely isolated. In the plant beds of St. John, about 9,000 feet in thickness, and in the coal formation of the South Joggins, more than 7,000 feet in thickness, no other Gasteropods occur, nor, I believe, do any occur in the beds holding land snails in Illinois. Nor, as already stated, are any of the aquatic Pulmonifera known in the Palæozoic. Thus, in so far as at present known, these Palæozoic snails are separated not only from any predecessors, if there were any, or successors, but from any contemporary animals allied to them.

It is probable that the land snails of the Erian and Carboniferous were neither numerous nor important members of the faunas of those periods. Had other species existed in any considerable numbers, there is no reason why they should not have been found in the erect trees, or in those shales which contain land plants. More especially would the discovery of any larger species, had they existed, been likely to have occurred. Further, what we know of the vegetation of the Palæozoic period would lead us to infer that it did not aboundin those succulent and nutritious leaves and fruits which are most congenial to land snails. It is to be observed, however, that we know little as yet of the upland life of the Erian or Carboniferous. The animal life of the drier parts of the low country is indeed as yet very little known; and but for the revelations in this respect of the erect trees in one bed in the coal formation of Nova Scotia, our knowledge of the land snails and Millipedes, and also of an eminently terrestrial group of reptiles, theMicrosauria, would have been much more imperfect than it is. We may hope for still further revelations of this kind, and in the meantime it would be premature to speculate as to the affinities of our little group of land snails with animals either their contemporaries or belonging to earlier or later formations, except to note the fact of the little change of form or structure in this type of life in that vast interval of time which separates the Erian period from the present day.

It may be proper to mention here the alleged Pulmonifera of the genusPalæorbisdescribed by some German naturalists. These I believe to be worm tubes of the genusSpirorbis, and in fact to be nothing else than the commonS. carbonariusorS. pusillusof the coal formation. The history of this error may be stated thus. The eminent palæobotanists Germar, Goeppert and Geinitz have referred theSpirorbis, so common in the Coal measures to the fungi, under the nameGyromyces, and in this they have been followed by other naturalists, though as long ago as 1868 I had shown that this little organism is not only a calcareous shell, attached by one side to vegetable matters and shells of mollusks, but that it has the microscopic structure characteristic of modern shells of this type.[139]More recently Van Beneden, Cænius, and Goldenberg, perceiving that the fossil is really a calcareous shell, butapparently unaware of the observations made in this country by myself and Mr. Lesquereux, have held theSpirorbisto be a pulmonate mollusk allied toPlanorbis, and have supposed that its presence on fossil plants is confirmatory of this view, though the shells are attached by a flattened side to these plants, and are also found attached to shells of bivalves of the genusNaiadites. Mr. R. Etheridge, jun., of the Geological Survey of Great Britain, has summed up the evidence as to the true nature of these probably brackish-water shells, and has revised and added to the species, in a series of articles in theGeological Magazineof London, vol. viii.

[139]"Acadian Geology," 2nd edition, p. 205.

[139]"Acadian Geology," 2nd edition, p. 205.

The erect trees of Coal Mine Point are rich in remains of Millipedes. The first of these (Xylobius Sigillariæ), which was the first known Palæozoic Myriapod, was described by me from specimens found in a tree extracted in 1852, and this, with a number of other remains subsequently found, was afterwards placed in the hands of Dr. Scudder, who has recognised in the material submitted to him eight species belonging to three genera (Xylobius,Archiulus, andAmynilyspes). These animals in all probability haunted these trees to feed on the decaying wood and other vegetable matter, and were undoubtedly themselves the prey of the Microsaurians. Though these were the earliest known, their discovery was followed by that of many other species in Europe and America, and some of them as old as the Devonian.[140]

[140]The two first-named genera from the erect trees, according to Scudder, belong to an extinct family of Millipedes, which he names Archiulidæ, and places with other Carboniferous genera in the orderArchipolypoda. The third belongs to family Euphoberidæ. Proc. R. S. of London, 1892.

[140]The two first-named genera from the erect trees, according to Scudder, belong to an extinct family of Millipedes, which he names Archiulidæ, and places with other Carboniferous genera in the orderArchipolypoda. The third belongs to family Euphoberidæ. Proc. R. S. of London, 1892.

The only other remains of Air-breathers found in the erect trees belong to Scorpions, of which some fragments remain in such a state as to make it probable that they have been partially devoured by the imprisoned reptiles. No remains of any aquatic animals have been found in these trees. TheScorpions are referred by Scudder to three species belonging to two genera.[141]

[141]Mazonia Acadica, and a second species ofMazonia, with fragments of a third species, generally distinct. Proceedings Royal Society of London, 1892.

[141]Mazonia Acadica, and a second species ofMazonia, with fragments of a third species, generally distinct. Proceedings Royal Society of London, 1892.

In the previous paper we have considered the mode of accumulation of Coal, and it may be useful here to note the light thrown on this subject by the Air-breathers of the coal formation and their mode of occurrence.

In no part of the world are the coal measures better developed, or more fully exposed, than in the coast sections of Nova Scotia and Cape Breton; and in these, throughout their whole thickness, no indication has been found of any of the marine fossils of the Lower Carboniferous Limestone. Abundant remains of fishes occur, but these may have frequented estuaries, streams and ponds, and the greater part of them are small ganoids which, like the modernLepidosteusandAmia, may have been specially fitted by their semi-reptilian respiration, for the impure waters of swampy districts. Bivalve mollusks also abound; but these are all of the kinds to which I have given the generic nameNaiadites, and Mr. Salter those ofAnthracomyaandAnthracoptera. These shells are all distinct from any known in the marine limestones. Their thin edentulous valves, their structure consisting of a wrinkled epidermis, a thin layer of prismatic shell and an inner layer of imperfectly pearly shell, all remind us of the Anodons and Unios. A slight notch in front concurs with their mode of occurrence in rendering it probable that, like mussels in modern estuaries, they attached themselves to floating or sunken timber. They are thus removed, both in structure and habit, from truly marine species; and may have been fresh-water or brackish-water mussels closely allied to modern Unios.

Carboniferous Millipedes,Xylobius Sigillariæ, Dawson (a, c), andArchiulus xylobioides, Scudder (b).Carboniferous Cockroach.—Blattina Bretonensis, Sc.Carboniferous Scorpion.—Anthracomartus Carbonarius, abdominal segments.

Carboniferous Millipedes,Xylobius Sigillariæ, Dawson (a, c), andArchiulus xylobioides, Scudder (b).

The crustaceans (Eurypterus,Diplostylus,Cyprids), and the worm shell (Spirorbis) found with them, are not necessarily marine, though some of them belonged probably to brackish water, and they have not yet been found in those carboniferous beds deposited in the open sea. There is thus in the whole thickness of the middle coal measures of Nova Scotia a remarkable absence at least of open sea animals; and if, as is quite probable, the sea inundated at intervals the areas of coal accumulation, the waters must have been shallow, and to a great extent land-locked, so that brackish-water rather than marine animals inhabited them.

On the other hand, there are in these coal measures abundant evidences of land surfaces; and subaërial decay of vegetable matter in large quantity is proved by the occurrence of the mineral charcoal of the coal itself, as I have elsewhere shown.[142]The erect trees which occur at so many levels also imply subaërial decay. A tree imbedded in sediment and remaining under water, could not decay so as to become hollow and deposit the remains of its wood in the state of mineral charcoal within the hollow bark. Yet this is the case with the greater part of the erect Sigillariæ which occur at more than twenty levels in the Joggins section. Nor could such hollow trunks become repositories for millipedes, snails and reptiles, if under water. On the other hand, if, as seems necessary to explain the character of the reptiliferous erect trees, these remained dry, or nearly so, in the interior, this would imply not merely a soil out of water, but comparatively well drained; as would indeed always be the case, when a flat resting on a sandy subsoil was raised several feet above the level of the water. Further, though the peculiar character of the roots ofSigillariæandCalamitesmay lend some countenance to the supposition that they could grow under water, or in water-soaked soils, this will not apply to coniferous trees, toferns, and other plants, which are found under circumstances which show that they grew with theSigillariæ.

[142]Journal of Geological Survey, vol. xv.

[142]Journal of Geological Survey, vol. xv.

In the coal measures of Nova Scotia, therefore, while marine conditions are absent, there are ample evidences of fresh-water or brackish-water conditions, and of land surfaces, suitable for the air-breathing animals of the period. Nor do I believe that the coal measures of Nova Scotia were exceptional in this respect. It is true that in Great Britain evidences of marine life do occur in the coal measures; but not, so far as I am aware, in circumstances which justify the inference that the coal is of marine origin. Alternations of marine and land remains, and even mixtures of these, are frequent in modern submarine forests. When we find, as at Fort Lawrence in Nova Scotia, a modern forest rooted in upland soil forty feet below high-water mark,[143]and covered with mud containing living Tellinas and Myas, we are not justified in inferring that this forest grew in the sea. We rather infer that subsidence has occurred. In modern salt marshes it is not unusual to find every little runnel or pool full of marine shell fish, while in the higher parts of the marsh land plants are growing; and in such places the deposit formed must contain a mixture of land plants and marine animals with salt grasses and herbage—the wholein situ.[144]

[143]Journal of Geological Society, vol. xi.[144]In the marshes at the mouth of Scarborough River, in Maine, channels not more than a foot wide, and far from the sea, are full of Mussels and Myæ; and in little pools communicating with these channels there are often many youngLimuli, which seem to prefer such places, and the cast-off shells and other remains of which may become imbedded in mud and mixed with land plants, just as in the shales of the coal measures.

[143]Journal of Geological Society, vol. xi.

[144]In the marshes at the mouth of Scarborough River, in Maine, channels not more than a foot wide, and far from the sea, are full of Mussels and Myæ; and in little pools communicating with these channels there are often many youngLimuli, which seem to prefer such places, and the cast-off shells and other remains of which may become imbedded in mud and mixed with land plants, just as in the shales of the coal measures.

These considerations serve, I think, to explain all the apparently anomalous associations of coal plants with marine fossils; and I do not know any other arguments of apparent weight that can be adduced in favour of the marine or evenaquatic origin of coal, except such as are based on misconceptions of the structure and mode of growth of sigillaroid trees and of the stratigraphical relations of the coal itself.[145]It is to be observed, however, that while I must maintain the essentially terrestrial character of the ordinary coal and of its plants, I have elsewhere admitted that cannel coals and earthy bitumen present evidences of subaquatic deposition; and have also abundantly illustrated the facts that the coal plants grew on swampy flats, liable not only to river inundations, but also to subsidence and submergence.[146]In the oscillation of these conditions it is evident thatSigillariæand their contemporaries must often have been placed in conditions unfavourable or fatal to them, and when their remains are preserved to us in these conditions, we may form very incorrect inferences as to their mode of life. Further, it is to be observed that the conditions of submergence and silting up which were favourable to the preservation of specimens ofSigillariæas fossils, must have been precisely those whichwere destructive to them as living plants; and on the contrary, that the conditions in which these forests may have flourished for centuries must have been those in which there was little chance of their remains being preserved to us, in any other condition at least than that of coal, which reveals only to careful microscopic examination the circumstances, whether aërial or aquatic, under which it was formed.

[145]It is unfortunate that few writers on this subject have combined with the knowledge of the geological features of the coal a sufficient acquaintance with the phenomena of modern marshes and swamps, and with the conditions necessary for the growth of plants such as those of the coal. It would be easy to show, were this a proper place to do so, that the "swells," "rock faults," splitting of beds, and other appearances of coal seams quite accord with the theory of swamp accumulation; that the plants associated withSigillariæcould not have lived with their roots immersed in salt water; that the chemical character of the underclays implies drainage and other conditions impossible under the sea; that the composition and minute structure of the coal are incompatible with the supposition that it is a deposit from water, and especially from salt water; and that it would be more natural to invoke wind driftage as a mode of accumulation for some of the sandstones, than water driftage for the formation of the coal. At the same time it is pretty certain that such beds as the cannels and earthy bitumens which appear to consist of finely comminuted vegetable matter, without mineral charcoal, may have been deposits of muck in shallow lakes or lagoons.[146]Journal of Geol. Socy., vols. x. and xv., and "Acadian Geology."

[145]It is unfortunate that few writers on this subject have combined with the knowledge of the geological features of the coal a sufficient acquaintance with the phenomena of modern marshes and swamps, and with the conditions necessary for the growth of plants such as those of the coal. It would be easy to show, were this a proper place to do so, that the "swells," "rock faults," splitting of beds, and other appearances of coal seams quite accord with the theory of swamp accumulation; that the plants associated withSigillariæcould not have lived with their roots immersed in salt water; that the chemical character of the underclays implies drainage and other conditions impossible under the sea; that the composition and minute structure of the coal are incompatible with the supposition that it is a deposit from water, and especially from salt water; and that it would be more natural to invoke wind driftage as a mode of accumulation for some of the sandstones, than water driftage for the formation of the coal. At the same time it is pretty certain that such beds as the cannels and earthy bitumens which appear to consist of finely comminuted vegetable matter, without mineral charcoal, may have been deposits of muck in shallow lakes or lagoons.

[146]Journal of Geol. Socy., vols. x. and xv., and "Acadian Geology."

It is also noticeable that, in conditions such as those of the coal formation, it would be likely that some plants would be specially adapted to occupy newly emerged flats and places liable to inundation and silting up. I believe that many of the Sigillariæ, and still more eminently theCalamites, were suitable to such stations. There is direct evidence that the nuts named Trigonocarpa were drifted extensively by water over submerged flats of mud. ManyCardiocarpawere winged seeds which may have drifted in the air. The Calamites may, like modernEquiseta, have produced spores with elaters capable of floating them in the wind. One of the thinner coals at the Joggins is filled with spores or spore cases that seem to have carried hairs on their surfaces, and may have been suited to such a mode of dissemination. I have elsewhere proved[147]that at least some species ofCalamiteswere, by their mode of growth, admirably fitted for growing amid accumulating sediment, and for promoting its accumulation.

[147]"Acadian Geology," chapter on Coal Plants.

[147]"Acadian Geology," chapter on Coal Plants.

The reptiles of the coal formation are probably the oldest known to us, and possibly, though this we cannot affirm, the highest products of creation in this period. Supposing, for the moment, that they are the highest animals of their time, and, what is perhaps less likely, that those which we know are a fair average of the rest, we have the curious fact that they are all carnivorous, and the greater part of them fitted to find food in the water as well as on the land. The plant feeders of the period, on the land at least, are all invertebrates, as snails,millipedes, and perhaps insects. The air-breathing vertebrates are not intended to consume the exuberant vegetable growth, but to check the increase of its animal enemies. Plant life would thus seem to have had in every way the advantage. The millipedes probably fed only on roots and decaying substances, the snails on the more juicy and succulent plants growing in the shadow of the woods, and the great predominance of the family of cockroaches among carboniferous insects points to similar conclusions as to that class. While, moreover, the vegetation of the coal swamps was most abundant, it was not, on the whole, of a character to lead us to suppose that it supported many animals. Our knowledge of the flora of the coal swamps is sufficiently complete to exclude from them any abundance of the higher phænogamous plants. We know little, it is true, of the flora of the uplands of the period; but when we speak of the coal-formation land, it is to the flats only that we refer. The foliage of the plants on these flats with the exception of that of the ferns, was harsh and meagre, and there seem to have been no grasses or other nutritious herbaceous plants. These are wants of themselves likely to exclude many of the higher forms of herbivorous life. On the other hand, there was a profusion of large nut-like seeds, which in a modern forest would probably have afforded subsistence to squirrels and similar animals. The pith and thick soft bark of many of the trees must at certain seasons have contained much nutritive matter, while there was certainly sufficient material for all those insects whose larvæ feed on living and dead timber, as well as for the creatures that in turn prey on them. It is remarkable that there seem to have been no vertebrate animals fitted to avail themselves of these vast stores of food. The question: "What may have fed on all this vegetation?" was never absent from my mind in all my explorations of the Nova Scotia coal sections; but no trace of any creature other than those already mentioned has ever rewarded my search. InNova Scotia it would seem that a few snails, gally-worms, and insects were the sole links of connection between the plant creation and air-breathing vertebrates. Is this due to the paucity of the fauna, or the imperfection of the record? The fact that a few erect stumps have revealed nearly all the air-breathers yet found, argues strongly for the latter cause; but there are some facts bearing on the other side.

A gally-worm, if, like its modern relatives, hiding in crevices of wood in forests, was one of the least likely animals to be found in aqueous deposits. The erect trees gave it its almost sole chance of preservation.Pupa vetustais a small species, and its shell very thin and fragile, while it probably lived among thick vegetation. Further, the measures 2,000 feet thick, separating the lowest and highest beds in which it occurs, include twenty-one coal seams, having an aggregate thickness of about twenty feet, three beds of bituminous limestone of animal origin, and perhaps twenty beds holdingStigmariain situ, orerectSigillariæandCalamites. The lapse of time implied by this succession of beds, many of them necessarily of very slow deposition, must be very great, though it would be mere guess work to attempt to resolve it into years. Yet long though this interval must have been,Pupa vetustalasted without one iota of change through it all; and, more remarkable still, was not accompanied by more than two other species of its family. Where so many specimens occur, and in situations so diverse, without any additional species, the inference is strong that no other of similar habits existed. If in any of those subtropical islands, whose climate and productions somewhat resemble those of the coal period, after searching in and about decaying trees, and also on the bars upon which rivers and lakes drifted their burdens of shells, we should find only three species, but one of these in very great numbers, we would surely conclude that other species, if present, were very rare.

Again, footprints referable toDendrerpeton, or similar animals,occur in the lower Carboniferous beds below the marine limestones, in the middle coal measures, and in the upper coal formation, separated by a thickness of beds which may be estimated at 15,000 feet, and certainly representing a vast lapse of time. Did we know the creature by these impressions alone, we might infer its continued existence for all this great length of time; but when we also find its bones in the principal repositories of reptile remains, and in company with the other creatures found with it, we satisfy ourselves that of them all it was the most likely to have left its trail in the mud flats. We thus have reason to conclude that it existed alone during this period, in so far as its especial kind of habitat was concerned; though there lived with it other reptiles, some of which, haunting principally the woods, and others the water, were less likely to leave impressions of their footprints. These may be but slight indications of truth, but they convey strong impressions of the persistence of species, and also of the paucity of species belonging to these tribes at the time.

If we could affirm that the Air-breathers of the coal period were really the first species of their families, they might acquire additional interest by their bearing on this question of origin of species. We cannot affirm this; but it may be a harmless and not uninstructive play of fancy to suppose for a moment that they actually are so, and to inquire on this supposition as to the mode of their introduction. Looking at them from this point of view, we shall first be struck with the fact that they belong to all of the three great leading types of animals which include our modern Air-breathers the Vertebrates, the Arthropods, and the Mollusks. We have besides to consider in this connection that the breathing organs of an insect are air tubes opening laterally (tracheæ), those of a land snail merely a modification of the chamber which in marine species holds the gills, while those of the reptiles represent the air bladder of the fishes. Thus, in the three groups the breathing organs arequite distinct in their nature and affinities. This at once excludes the supposition that they can all have been derived from each other within the limits of the coal period. No transmutationist can have the hardihood to assert the convertibility, by any direct method, of a snail into a millipede or an insect, or of either into a reptile. The plan of structure in these creatures is not only different, but contrasted in its most essential features. It would be far more natural to suppose that these animals sprang from aquatic species of their respective types. We should then seek for the ancestors of the snail in aquatic Gasteropods, for those of the millipede in worms or Crustaceans, and for those of the reptiles in the fishes of the period. It would be easy to build up an imaginary series of stages, on the principle of natural selection, whereby these results might be effected; but the hypothesis would be destitute of any support from fact, and would be beset by more difficulties than it removes. Why should the result of the transformation of water-snails breathing by gills be aPupa ?Would it not much more likely be anAuriculaor aLimnea ?It will not solve this difficulty to say that the intermediate forms became extinct, and so are lost. On the contrary, they exist to this day, though they were not, in so far as we know, introduced so early. But negative evidence must not be relied on; the record is very imperfect, and such creatures may have existed, though unknown to us. It may be answered that they could not have existed in any considerable numbers, else some of their shells would have appeared in the coal-formation beds, so rich in crustaceans and bivalve mollusks. Further, the little Pupa remained unchanged during a very long time, and shows no tendency to resolve itself into anything higher, or to descend to anything lower, while in the lowest bed in which it occurs it is associated with a round snail of quite different type. Here, if anywhere, in what appears to be the first introduction of air-breathing invertebrates, we should be able to find theevidences of transition from the gills of the Prosobranchiate and the Crustacean to the air sac of the Pulmonate and the tracheæ of the millipede. It is also to be observed that many other structural changes are involved, the aggregate of which makes a Pulmonate or a millipede different in every particular from its nearest allies among gill-bearing Gasteropods or Crustaceans.

It may be said, however, that the links of connection between the coal reptiles and fishes are better established. All the known coal reptiles have leanings to the fishes in certain characters; and in some, as inArchegosaurus, these are very close. Still the interval to be bridged over is wide, and the differences are by no means those which we should expect. Were the problem given to convert a ganoid fish into anArchegosaurusorDendrerpeton, we should be disposed to retain unchanged such characters as would be suited to the new habits of the creature, and to change only those directly related to the objects in view. We should probably give little attention to differences in the arrangement of skull bones, in the parts of the vertebræ, in the external clothing, in the microscopic structure of the bone, and other peculiarities for serving similar purposes by organs on a different plan, which are so conspicuous so soon as we pass from the fish to the Batrachian. It is not, in short, an improvement of the organs of the fish that we witness so much as the introduction of new organs.[148]The foot of the batrachian bears, perhaps, as close a relation to the fin of the fish as the screw of one steamship to the paddle wheel of another, or as the latter to a carriage wheel; and can be just as rationally supposed to be not a new instrument, but the old one changed. In this connection even a footprint in the sand startles us as much as that of Friday did RobinsonCrusoe. We see five fingers and toes, and ask how this numerical arrangement started at once from fin rays of fishes all over the world; and how it has continued unchanged till now, when it forms the basis of our decimal arithmetic.

[148]An ingenious attempt by Prof. Cope, to deduce the batrachian foot from the fins of certain carboniferous fishes, will be found in theProceedings of the Philos. Academy of Philadelphiafor the present year.

[148]An ingenious attempt by Prof. Cope, to deduce the batrachian foot from the fins of certain carboniferous fishes, will be found in theProceedings of the Philos. Academy of Philadelphiafor the present year.

Again, our reptiles of the coal do not constitute a continuous series, and belong to a great number of distinct genera and families, nor is it possible that they can all, except at widely different times, have originated from the same source. It either happened, for some unknown reason, that many kinds of fishes put on the reptilian guise in the same period, or else the vast lapse of ages required for the production of a reptile from a fish must be indefinitely increased for the production of many dissimilar reptiles from each other; or, on the other hand, we must suppose that the limit between the fish and reptile being once overpassed, a facility for comparatively rapid changes became the property of the latter. Either supposition would, I think, contradict such facts bearing on the subject as are known to us.

We commenced with supposing that the reptiles of the Coal might possibly be the first of their family, but it is evident from the above considerations, that on the doctrine of natural selection, the number and variety of reptiles in this period would imply that their predecessors in this form must have existed from a time as early as any in which even fishes are known to exist; so that if we adopt any hypothesis of derivation, it would probably be necessary to have recourse to that which supposes at particular periods a sudden and as yet unaccountable transmutation of one form into another; a view which, in its remoteness from anything included under ordinary natural laws, does not materially differ from that currently received idea of creative intervention, with which, in so far as our coal reptiles can inform us, we are for the present satisfied.

There is one other point which strikes the naturalist in considering these animals, and which has a certain bearing on suchhypothesis. It is the combination of various grades of reptilian types in these ancient creatures. It has been well remarked by Hugh Miller, and more fully by Agassiz, that this is characteristic of the first appearance of new groups of animals. Now selection, as it acts in the hands of the breeder, tends to specialization; and natural selection, if there is such a thing, is supposed to tend in the same direction. But when some distinctly new form is to be introduced, an opposite tendency seems to prevail, a sort of aggregation in one species of characters afterward to be separated and manifested in distinct groups of creatures. The introduction of such new types also tends to degrade and deprive of their higher properties previously existing groups of lower rank. It is easy to perceive in all this, law and order, in that higher sense in which these terms express the will and plan of the Supreme Mind, but not in that lower sense in which they represent the insensate operation of blind natural forces.

References:—"Air-breathers of the Coal Period." Montreal, 1886. Papers on Reptiles, etc., in South Joggins Coal Field,Journal of Geological Society of London, vols. ix. x. xi. xvi. Remains of Animals in Erect Trees in the Coal Formation of Nova Scotia,Trans. Royal Society, 1881. "Acadian Geology," fourth edition, 1891. Revision of Land Snails of the Palæozoic Era,Am. Journal of Science, vol. xx., 1880. Supplementary Report to Royal Society of London, Proceedings, 1892. Notice of additional Reptilian Remains,Geological Magazine of London, 1891.

DEDICATED TO THE MEMORY OF THE LATEDR. J. J. BIGSBY, F.R.S.,OF LONDON,The painstaking and accurate Authorof the Thesaurus Siluricus and Devonico-Carboniferous,a warm and kind Friend and Christian Gentlemanand one of thePioneers of Canadian Geology.

Reminiscences of Lyell's Work—Tidal Flats of the Bay of Fundy—Rill Marks and Shrinkage Cracks—Worm Trails and Burrows—The Paces of Limulus—Fucoids Versus Trails—Footprints of Vertebrates

Track of Limulus.—Modern, Orchard Beach. Showing its resemblance to theProtichnitesof the Cambrian. (Page 320.)

Track of Limulus.—Modern, Orchard Beach. Showing its resemblance to theProtichnitesof the Cambrian. (Page 320.)

MARKINGS, FOOTPRINTS AND FUCOIDS.

I

Ibelieve my attention was first directed to the markings made by animals on the surfaces of rocks, when travelling with the late Sir Charles Lyell in Nova Scotia, in 1842. He noticed with the greatest interest the trails of worms, insects, and various other creatures, and the footprints of birds on the surface of the soft red tidal mud of the Bay of Fundy, and subsequently published his notes on the various markings in these deposits in his "Travels in North America," and in a paper presented to the Geological Society of London. I well remember how, in walking along the edge of the muddy shore, he stopped to watch the efforts of a grasshopper that had leaped into the soft ooze, and was painfully making a most complicated trail in his effort to escape. Sir Charles remarked that if it had been so fortunate as to make these strange and complicated tracks on some old formation now hardened into stone and buried in the earth, it might have given occasion to much learned discussion.

At a later period I found myself perplexed in the study of fossil plants by the evident errors of many palæobotanists unacquainted with modern markings on shores, in referring all kinds of mere markings to the vegetable kingdom, and especially to the group of fucoids or seaweeds, which had become a refuge for destitute objects not referable to other kinds of fossils. It thus became necessary to collect and study these objects, as they existed in rocks of different ages, and to comparethem with the examples afforded by the modern beach; and perhaps no locality could have afforded better opportunities for this than the immense tidal flats of the finest mud left bare by the great tides of the Bay of Fundy in Nova Scotia. At a more recent period still, the subject has come into great prominence in Europe, and if we are to gauge its importance by the magnitude of the costly illustrated works devoted to it by Delgado, Saporta, Nathorst, and others, and the multitude of scattered papers in scientific periodicals, we should regard it as one of the most salient points in Geology.[149]

[149]Journal of London Geological Society, vol. vii. p. 239.

[149]Journal of London Geological Society, vol. vii. p. 239.

It may be well further to introduce the subject by a few extracts from Lyell's work above referred to.

"The sediment with which the waters are charged is extremely fine, being derived from the destruction of cliffs of red sandstone and shale, belonging chiefly to the coal measures. On the borders of even the smallest estuaries communicating with a bay, in which the tides rise sixty feet and upwards, large areas are laid dry for nearly a fortnight between the spring and neap tides, and the mud is then baked in summer by a hot sun, so that it becomes solidified and traversed by cracks caused by shrinkage. Portions of the hardened mud may then be taken up and removed without injury. On examining the edges of each slab we observe numerous layers, formed by successive tides, usually very thin, sometimes only one-tenth of an inch thick, of unequal thickness, however, because, according to Dr. Webster, the night tides rising a foot higher than the day tides throw down more sediment. When a shower of rain falls, the highest portion of the mud-covered flat is usually too hard to receive any impressions; while that recently uncovered by the tide, near the water's edge, is too soft. Between these areas a zone occurs almost as smooth and even as a looking-glass, on which every drop forms a cavity of circular or oval form; and if the shower betransient, these pits retain their shape permanently, being dried by the sun, and being then too firm to be effaced by the action of the succeeding tide, which deposits upon them a new layer of mud. Hence we find, on splitting open a slab an inch or more thick, on the upper surface of which the marks of recent rain occur, that an inferior layer, deposited perhaps ten or fourteen tides previously, exhibits on its under surface perfect casts of rain prints which stand out in relief, the moulds of the same being seen in the layer below."

After mentioning that a continued shower of rain obliterates the more regular impressions, and produces merely a blistered or uneven surface, and describing minutely the characteristics of true rain marks in their most perfect state, Sir Charles adds:—

"On some of the specimens the winding tubular tracks of worms are seen, which have been bored just beneath the surface. Sometimes the worms have dived beneath the surface, and then re-appeared. Occasionally the same mud is traversed by the footprints of birds (Tringa minuta), and of musk-rats, minks, dogs, sheep and cats. The leaves also of the elm, maple and oak trees have been scattered by the winds over the soft mud, and having been buried under the deposits of succeeding tides, are found on dividing the layers. When the leaves themselves are removed, very faithful impressions, not only of their outline, but of their minutest veins, are left imprinted on the clay."

This is a minor illustration of that application of recent causes to explain ancient effects of which the great English geologist was the apostle and advocate, and which he so admirably practised in his own work. It is also an illustration of the fact that things the most perishable and evanescent may, when buried in the crust of the earth, become its most durable monuments. Footprints in the sand of the tidal shore are in the ordinary course of events certain to be obliteratedby the next tide; but when carefully filled up by gently deposited new material, and hardened into stone, there is no limit to their duration.

Let us inquire how this may take place, and the tidal flats of the Bay of Fundy and Basin of Minas may supply us with the information desired. In the upper parts of the Bay of Fundy and its estuaries the rise and fall of tide, as is well known, are excessive. I quote the following description of the appearance they present from a work of earlier date:—

"The tide wave that sweeps to the north-east, along the Atlantic coast of the United States, entering the funnel-like mouth of the Bay of Fundy, becomes compressed and elevated, as the sides of the bay gradually approach each other, until in the narrower parts the water runs at the rate of six or seven miles per hour, and the vertical rise of the tide amounts to sixty feet or more. In Cobequid and Chiegnecto Bays these tides, to an unaccustomed spectator, have rather the aspect of some rare convulsion of nature than of an ordinary daily phenomenon. At low tide wide flats of brown mud are seen to extend for miles, as if the sea had altogether retired from its bed; and the distant channel appears as a mere strip of muddy water. At the commencement of flood a slight ripple is seen to break over the edge of the flats. It rushes swiftly forward, and, covering the lower flats almost instantaneously, gains rapidly on the higher swells of mud, which appear as if they were being dissolved in the turbid waters. At the same time the torrent of red water enters all the channels, creeks and estuaries; surging, whirling, and foaming, and often having in its front a white, breaking wave, or 'bore,' which runs steadily forward, meeting and swallowing up the remains of the ebb still trickling down the channels. The mud flats are soon covered; and then, as the stranger sees the water gaining with noiseless and steady rapidity on the steep sides of banks and cliffs, a sense of insecurity creeps over him, as if no limitcould be set to the advancing deluge. In a little time, however, he sees that the fiat, 'Hitherto shalt thou come, and no farther,' has been issued to the great bay tide: its retreat commences, and the waters rush back as rapidly as they entered.

"The rising tide sweeps away the fine material from every exposed bank and cliff, and becomes loaded with mud and extremely fine sand, which, as it stagnates at high water, it deposits in a thin layer on the surface of the flats. This layer, which may vary in thickness from a quarter of an inch to a quarter of a line, is coarser and thicker at the outer edge of the flats than nearer the shore; and hence these flats, as well as the marshes, are usually higher near the channels than at their inner edge. From the same cause,—the more rapid deposition of the coarser sediment,—the lower side of each layer is arenaceous, and sometimes dotted over with films of mica, while the upper side is fine and slimy, and when dry has a shining and polished surface. The falling tide has little effect on these deposits, and hence the gradual growth of the flats, until they reach such a height that they can be overflowed only by the high spring tides. They then become natural or salt marsh, covered with the coarse grasses andcariceswhich grow in such places. So far the process is carried on by the hand of nature; and before the colonization of Nova Scotia, there were large tracts of this grassy alluvium to excite the wonder and delight of the first settlers on the shores of the Bay of Fundy. Man, however, carries the land-making process farther; and by diking and draining, excludes the sea water, and produces a soil capable of yielding for an indefinite period, without manure, the most valuable cultivated grains and grasses."

The mud of these great tidal flats is at the surface of a red colour, and so fine that when the tide leaves it and its surface becomes dry, it shines in the sun as if polished. It is thus capable of taking the finest impressions. When the tide is in,numerous small fish of various species occupy the ground and may leave marks of their fins and tails as they gambol or seek their food. Shell fishes, worms, and Crustaceans scramble over the same surface, or make burrows in it. As the tide recedes flocks of sandpipers and crows follow it down, and leave an infinity of footprints, and even quadrupeds like the domestic hog go far out at low water in search of food. It is said that in some parts of the Bay the hogs are so assiduous in this pursuit that they even awake and go out on the flats in the night tide, and that they have so learned to dread the dangers of the flood, that when in the darkness they hear the dull sound of the approaching bore, they squeal with fear and rush madly for the shore.

If we examine it minutely, we shall find that the tidal deposit is laminated. The tidal water is red and muddy, and holds in suspension sediment of various degrees of coarseness. This, undergoes a certain process of levigation. In the first run of the flood the coarser material falls to the bottom. As its force diminishes the finer material is deposited, and at full tide, when the current has ceased, the finest of all settles, forming a delicate coat of the purest and most tenacious clay. Thus, if a block of the material is taken up and allowed to dry, it tends to separate into thin laminæ, each of which represents a tide, and is somewhat sandy below, and passes into the finest moulding clay above. The tracks and impressions preserved are naturally made on the last or finest deposit, and filled in with the coarser or more sandy of the next tide. But this may take place in different ways. Impressions made under water at flood tide, or on the surface left bare by the ebb, may in favourable localities be sufficiently tenacious or firm to resist the abrading action of the flood, and may thus be covered and preserved by the next layer, and in this way they may be seen on splitting up a block of the dried mud. But in shallow places and near the shore, where the deposithas time to consolidate and become dried by the sun and air before the next tide, much better impressions are preserved; and lastly, on those parts of the shore which are reached only by the spring tides, the mud of the highest tide of course may have several days to harden before the next tide reaches it, and in this case it becomes cracked by an infinity of shrinkage cracks, which, when it is next covered with the tide, are filled with new sediment. In this way is produced in great perfection that combination of footprints, or even of impressions of rain, with casts of cracks, which is so often seen in the older rocks. Where on the sides of channels or near the shore the mud has a considerable slope, another and very curious effect results. As the tide ebbs the water drains off the surface, or oozing out of the wet sand and mud, forms at the top of the bank minute grooves often no larger than fine threads. These coalesce and form small channels, and these, again, larger ones, till at low tide the whole sloping surface is seen to be covered with a smooth and beautiful tracery resembling the rivers on a map, or the impressions of the trunks and branches of trees, or the fronds of gigantic seaweeds. These "rill marks," as they have been called, are found in great abundance in the coal formation and triassic sandstones and shales, and I am sorry to say, have often been named and described as Fucoids, and illustrated by sumptuous plates. Sometimes these impressions are so fine as to resemble the venation of leaves, sometimes so large as to simulate trees, and I have even seen them complicated with shrinkage cracks, the edges of which were minutely crenulated by little rills running into them from the surface.

It is further to be noticed that all these markings and impressions on tidal shores may, when covered by succeeding deposits, appear either in intaglio or relief. On the upper surface they are of course sunken, but on the lower surface of the bed deposited on them they are in relief. It often happensalso that these casts in relief are the best preserved. This arises from the fact that the original moulds or impressions are usually made in clay, whereas the filling material is sandy, and the latter, infiltrated with calcareous or siliceous matter, may become a hard sandstone, while the clay may remain a comparatively soft shale. This tendency of casts rather than of moulds to be preserved sometimes produces puzzling effects. A cylindrical or branching trail thus often assumes the appearance of a stem, and any pits or marginal impressions assume the form of projections or leaves, and thus a trail of a worm or Gastropod or a rill mark may easily simulate a plant. It is to be observed, however, that these prominent casts are on the under side of the beds, that their material is continuous with that of the beds to which they belong, and that they are destitute of any carbonaceous matter. There are, however, cases where markings may be in relief, even on the upper surfaces of beds. The following are illustrations of this. Just as a man walking in newly fallen snow compresses it under his feet, and if the snow be afterwards drifted away or melted away by the sun, the compressed part resists longest, and may appear as a raised footmark, so tracks made on soft material may consolidate it so that if the soft mud be afterwards washed away the tracks may remain projecting. Again, worms eject earthy matter from their burrows, forming mounds, patches or raised ridges of various forms on the surface, and some animals burrow immediately under the surface, pushing up the mud over them into a ridge, while others pile up over their bodies pellets of clay, forming an archway or tunnel as they go. Zeiller has shown that the mole cricket forms curious roofed trails of this kind, and it seems certain that Crustaceans and marine worms of different kinds execute similar works, and that their roofed burrows, either entire or fallen in, produce curious imitations of branches of plants.

The great and multiform army of the sea worms is indeedthe most prolific source of markings on sea-formed rocks. Sometimes they cover very large surfaces of these, or penetrate the beds as perforations, with tortuous furrows, or holes perfectly simple, or marked with little striæ made by bristles or minute feet, sometimes with a fringe of little footmarks a each side, sometimes with transverse furrows indicating the joints of the animal's body. Multitudes of these markings have been described and named either as plants or as worm-tracks. Again, these creatures execute subterranean burrows, sometimes vertical, sometimes tortuous. These are often mere cylindrical holes afterwards filled with sand, but sometimes they have been lined with a membranous tube, or with the rejectamenta of the food of the animals, or with little fragments of organic matter cemented together. Sometimes they open on the surface as simple apertures, but again they may be surrounded with heaps of castings, sometimes spiral in form, or with dumps of sand produced in their excavation, and which may assume various forms, according to circumstances. Sometimes the aperture is double, so that they seem to be in pairs. Sometimes, for the convenience of the animal, the aperture is widened into the form of a funnel, and sometimes the creature, by extending its body and drawing it in, surrounds its burrow with a series of radiating tracks simulating the form of a starfish or sea anemone, or of the diverging branches of a plant.

Creatures of higher grade, provided with jointed limbs, naturally make their actions known in more complicated ways. Some years ago I had the pleasure of spending a few weeks at the favourite sea-side resort of Orchard Beach on the New England coast, and there made my first acquaintance with that very ancient and curious creature the Limulus, or Horse-shoe Crab, or King-crab, as it is sometimes called. Orchard Beach is, I presume, near its northern range on our coast, and the specimens seen were not very large in size, though by no meansrare, and not infrequently cast on shore in storms. But the best facilities for studying their habits were found in a marsh at no great distance from the hotel, where there were numerous channels, ditches and little ponds filled with sea water at high tide. In these were multitudes of young Limuli, varying from an inch to three or four inches in breadth, and though many were dead or merely cast shells, it was easy to take young specimens with a landing net. A number of these were secured, and I made it my business for some time to study their habits and mode of life, and especially the tracks which they made in sand or mud.

The King-crab, viewed from above, consists of three parts. The anterior shield or carapace is semi-circular in form, with two spines or projecting points at the angles, raised in the middle and sloping down to a smooth or moderately sharp edge in front. The eyes are set like windows in this shield. Two large ones at the sides, which are compound eyes consisting of numerous ocelli or little eyes, and two microscopic ones in front, at the base of a little spine, which are simple. The second or abdominal part is also in one piece, somewhat quadrate in form, with ridges and serratures at the sides armed with spines, and which may be said to simulate the separate joints into which the abdomen of an ordinary Crustacean is divided. The third part is a long tail spine, triangular in cross section, sharply pointed, and so jointed to the posterior end of the abdomen that it can be freely moved in any direction as a bayonet-like weapon of defence. When unable to escape from an enemy it is the habit of the creature to double itself up by bending the abdomen against the carapace, and erecting the sharp spine. Thus, with fixed bayonet it awaits attack, like the kneeling soldier in front of a square.

Below this upper shield, which is thin and papery in the young, somewhat horny in the adult, are the numerous limbs of the creature, with which we are at present most concerned.Under the carapace are several pairs of jointed limbs differing in size and form. The two anterior are small and peculiarly formed claws, used apparently in manipulating the food. The four next are larger in size, and are walking feet, each furnished with two sharp points which form a pincer for holding. The last pair is much larger and stronger than any of the others, and armed not only with a pair of pincers, but with four blunt nail-like points. Under the abdomen are flat swimming feet, as they have been called, each composed of a broad plate notched and divided in the middle. When at rest these lie flat on each other, but they can be flapped back and forth at the will of the animal.

Let us now see what use the creature can make of these numerous and varied pedal appendages, and for distinctness' sake we shall call the anterior set thoracic and the posterior abdominal. When placed in shallow water on fine sand it walked slowly forward, and its tracks then consisted of a number of punctures on the sand in two lines. If, however, the water was very shallow or the sand very soft or inclined upward the two edges of the carapace touched the bottom, making a slight furrow at each side; and if the tail was trailed on the bottom, this made a third or central furrow. When climbing a slope, or when placed at the edge of the water, it adopted another mode of locomotion, pushing with great force with its two posterior limbs, and thus moving forward by jerks. It then made four deep marks with the toes of each hind limb, and more or less interrupted marks with the edges of the carapace and the tail. In these circumstances the marks were almost exactly like those of some forms of the Protichnites of the Potsdam sandstone. When in sufficiently deep-water and desirous to escape, it flapped its abdominal feet, and then swam or glided close to the bottom. In this case, when moving near the soft bottom, it produced a series of transverse ridges and furrows like small ripple marks, with a slight ridge in the middle,and sometimes, when the edges of the carapace touched the bottom, with lateral furrows. In this way the animals were able to swim with some ease and rapidity, and on one occasion I observed an individual, confined in a tub of water, raise itself from the bottom and swim around the tub at the surface in search of a way of escape. Lastly, the young Limuli were fond of hiding themselves by burrowing in the sand. They did this by pushing the anterior rounded end of the carapace under the sand, and then vigorously shovelling out the material from below with their feet, so that they gradually sank under the surface, and the sand flowed in upon them till they were entirely covered. If carefully removed from the hollow they had made, this was found to be ovoid or hoof shaped in form and bilobed, not unlike the curious hollows (Rusophycus Grenvillensisof Billings) which I have supposed to be burrows of Trilobites.

I thus found that the common King-crab could produce a considerable variety of tracks and burrows comparable with those which have been named Protichnites, Climactichnites, Bilobites, Cruziana, Rusichnites, etc.; and that the kind of markings depended partly on the differences of gait in the animal, and partly on the circumstances in which it was placed; so that different kinds of tracks do not always prove diversity in the animals producing them.

The interest of this investigation as applied to Limulus is increased by the fact that this creature is the near ally of Trilobites, Eurypterids and other Crustaceans which were abundant in the earlier geological ages, and whose footprints are probably among the most common we find on the rocks.


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