A. Geological Relations of Eozoon, Archæozoon, etc.

B. Preservation of Organic Remains by Injection with Hydrous Silicates.

These minerals approach in composition to the jollyte of Von Kobell, from which they differ in containing a portion of alkalies, and only one half as much water. In these respects they agree nearly with the silicate found by Robert Hoffman, at Raspenau, in Bohemia, where it occurs in thin layers alternating with picrosmine, and surrounding masses of Eozoon in the Laurentian limestones of that region;[51] the Eozoon itself being there injected with a hydrous silicate which may be described as intermediate between glauconite and chlorite in composition."

In the Welsh specimen the silicate is of a deep green colour, except where oxidized, and though only 3 per cent, of the whole, is sufficient to give it an olive colour and slight serpentinous lustre. In the Pole Hill material, the silicate amounts to 5 per cent, of the whole, and is of a greyish colour. For some further particulars, see my Paper on "Fossils Mineralized with Silicates" (Journal Geological Society, February, 1879).

C. Affinities of Eozoon, etc., with more Modern Forms.

Dr. Carpenter, who in admirable papers, which I need not quote here,[52] has illustrated in detail the structures of Eozoon, and shown its resemblance to modern forms, places Eozoon as a generalized type between the Nummuline and Rotaline groups of Foraminifera. It resembles the former in its fine and complicated tubulations, and some of the larger sessile forms of the latter in its habit of growth. More especially, this is near to that of the genera Carpenteria and Polytrema. In the former, more especially, there are a number of somewhat flattened calcareous cells with perforated walls, and built up in a conical form around a central pipe or funnel into which the apertures of the cells open. A specimen of Carpenteria, enlarged and having the walls of its cells thickened by a supplemental tubulated deposit like that of Calcarina, would approach very near to Eozoon.

The question of the general relation of an organism like Eozoon to creatures known to us in the modern seas may be answered in either of two ways:—(1) Functionally or in relation to the position of such an animal in nature: or (2) Zoologically, or with reference to its affinities to other animals. With reference to the first consideration, the answer is plain. The geological function of Eozoon was that of a collector of calcareous matter from the surrounding waters, then probably very rich in calcium carbonate, and its role was the same with that of the Stromatoporæ and calcareous Sponges, smaller Foraminifera and Corals in latter times. The answer to the second aspect of the question is less easy. An ordinary observer would at once place Eozoon with the Stromatoporidæ or Layer-corals, which fill or even constitute whole beds of limestone in the Cambro-Silurian, Silurian and Devonian Periods. While, however, Eozoon has been claimed on the highest authority for the Rhizopods, the Stromatoporæ and their allies have been regarded as Sponges, or more recently as Hydroids allied to the Hydractiniæ and Millepores.[53] I confess that I am not satisfied with these interpretations. I have in my collections large numbers of encrusting spinous forms, usually called Stromatoporæ, but which I have long set aside as probably Hydractiniæ. There are other forms with large vertical tubes which I have regarded as corals, but some Stromatoporæ seem to be different from either, and I am still disposed to regard many of them as Protozoa. Bearing in mind, however, that the Silurian is as remote from the Laurentian on the one hand as from the Tertiary on the other, we might be prepared to expect that if the Layer-corals of the Silurian are divisible into different groups, somewhat widely separated, and we have in the lower Palæozoic the peculiar type of Cryptozoon, we may be prepared to expect in the Laurentian much more generalized forms, less susceptible of classification in our modern systems. If, therefore, Eozoon were accessible to us in a living state, I should not be surprised to find that—while perhaps more akin to the calcareous-shelled Rhizopods than to any other modern group—it may have presented points of resemblance to Sponges or even to Hydroids, in its skeleton and mode of growth, and even in the arrangement of its soft parts.

Taking this view of its nature and relations, the genus and the Laurentian species may be characterized as follows:—

Genus Eozoon, Dawson.

Foraminiferal skeletons, with irregular and often confluent cells, arranged in concentric and horizontal laminæ, or sometimes piled in an acervuline manner. Septal orifices irregularly disposed. Proper wall finely tubulated. Intermediate skeleton with branching canals.

Eozoon Canadense, Dawson.

In inverted conical or rounded masses or thick encrusting sheets, frequently of large dimensions. Typical structure stromatoporoid, or with concentric calcareous walls, frequently uniting with each other, and separating flat chambers, more or less mammillated, and spreading into horizontal lobes and small chamberlets; chambers often confluent and crossed by irregular calcareous pillars connecting the opposite walls. Upper part often composed of acervuline chambers of rounded forms. Proper wall tubulated very finely. Intermediate skeleton largely developed, especially at the lower part, and traversed by large branching canals, often with smaller canals in their interstices. Lower laminæ and chambers often three millimetres in thickness. Upper laminæ and chambers one millimetre or less. Age Upper Laurentian and perhaps Huronian.

Var. minor.—Supplemental skeleton wanting, except near the base, and with very fine canals. Laminæ of sarcode much mammillated, thin, and separated by very thin walls. Probably a depauperated variety.

Var. acervulina.—In oval or rounded masses, wholly acervuline. Cells rounded; intermediate skeleton absent or much reduced; cell-walls tubulated. This may be a distinct species, but it closely resembles the acervuline parts of the ordinary form.

Assuming the Archæospherinæ so abundantly found in the Eozoon limestones to be distinct organisms, and not mere germs or buds of Eozoon, they may be thus defined:—

Genus Archæospherina, Dawson.

A provisional genus, to include rounded solitary chambers, or globigerine assemblages of such chambers, with the cell-wall surrounding them tubulated as in Eozoon, or perhaps in some cases with simple pores like those of Rotalines. They may be distinct organisms, or gemmæ, or detached fragments of Eozoon. Some of them much resemble the bodies figured by Dr. Carpenter, as gemmæ or ova and primitive chambers of Orbitolites. They are very abundant on some of the strata surfaces of the limestones at Côte St. Pierre. Age Upper Laurentian.

I may add here the characters of Matthew's new genus, Archæozoon, as given by him:—

Genus Archæospherina, Matthew.

Skeleton composed of thin concentric laminæ convex upward, and having between them a granular layer filled with minute branching canals.

Archæospherina Acadiense, Matthew.

Habit of growth cylindrical in masses or groups, budding upward. The microscopic characters are thus given by Matthew[54]:—

"The structures appear to be allied more closely to Cryptozoon than to Eozoon. The microscopic structure is most easily recognised in the earthy (as distinguished from the calcareous) layers, and consists of minute branching canals. Under a one-inch objective the smaller canals have the appearance of minute threads, which run sometimes for a distance of two millimetres without branching. The larger canals branch more frequently and are more sinuous. The canals cross and anastomose with each other; they run chiefly at right angles to the axis of the fossil, and appear to branch most in going outward from the centre. More rarely they ascend from the earthy to the calcareous layer, branching upward."

In limestone of the Upper Laurentian, near St. John, New Brunswick.

D. Cryptozoon.

Professor Hall having kindly presented some good specimens to the Peter Redpath Museum, I have had sections made, and have thus been able to verify his description, and to compare the structures with those of some of the more ancient Stromatoporoid specimens in our collections, including the Archæozoon from New Brunswick, of which Mr. Matthew has presented a fine slab to the Museum. I have also, through the kindness of Professor Winchell, been enabled to compare these with his Cryptozoon Minnesotense, and Dr. Walcott has added specimens of his Stromatoporoid forms from the pre-Cambrian beds of Arizona. It would appear from these and other specimens in our collections from the Cambrian and older Ordovician beds, that we have here an ancient type of Stromatoporoid organism in which the original laminæ seem to have been thin and coriaceous, without apparent pores or pillars connecting them with each other, but having between them relatively-thick layers of fine fragmental matter penetrated by numerous irregularly tortuous and branching tubes. The laminæ often present a carbonaceous or chitinous appearance, though frequently replaced by mineral matter, and the intervening layers show both a calcareous and carbonaceous substance, with much fine silicious sand often as rounded grains, and apparently some dolomitic granules. The tubules seem destitute of any distinct wall, otherwise the whole would resemble on a large scale the nodular and laminated masses of Girvanella, which Wethered has described as surrounding organic fragments in Silurian and Carboniferous and Jurassic limestones in England.[55]

The Streptochetus of Seely from the Chazy limestone[56] is evidently very near to Girvanella, if not generically identical, and I have a similar species from the Lower Cambrian pebbles in the conglomerates of the Quebec group. In all these forms, however, the thicker or intermediate laminæ seem to consist wholly of definite convoluted tubes, whereas in Cryptozoon the tubes, or tubular perforations, are separated by a mass of material which in the best preserved specimens seems to consist of a fibrous stroma including calcareous and silicious particles. It seems doubtful to what class of beings such a structure should be referred; but whatever its nature, it evidently had great powers of growth, and seems to be a very ancient form of life.

One of the species similar in structure to Hall's type, but budding out into turbinate branches, was discovered by Mr. E. T. Chambers, of Montreal, in the Ordovician limestone of Lake St. John, and has been named C. boreale. It differs in structure from Hall's species in having the tubes less tortuous and more nearly parallel to the laminæ. In its outline it reminds one of the problematical Eozoon from the Hastings group at Tudor, Ontario, referred to in the text.

Should time permit, I hope to have all the specimens in our collections illustrating this interesting and primitive type examined and described. In the meantime I may merely remark that a near modern analogue would seem to be the gigantic arenaceous Foraminifer Neusina Agassizi, Goës, dredged by Alexander Agassiz in the Pacific, and described in the Bulletin of the Museum of Comparative Zoology (Vol. xxiii., No. 5, 1892). The modern form, it is true, is flat and foliaceous; but some of the old species approach to this shape, and if we suppose the little cells of Neusina to represent the tubes of Cryptozoon, and the carbonaceous matter of the latter to be the remains of the chitinous stroma seen in some specimens, the general resemblance will be very close.

The whole subject of these peculiar Stromatoporoid forms extending from the Upper Cambrian to the Laurentian, deserves a full and careful investigation, for which I am endeavouring to collect material.

E. Receptaculites and Archæocyathus.

In "The Dawn of Life" (1875), reference was made to the singular and complicated organisms of the Upper Cambrian and Ordovician systems known as Receptaculites, which at that time was generally regarded as foraminiferal, and is still placed by Zittel, in his great work on Palæontology, among forms doubtfully referable to that group. It has also been referred to Sponges, though on very uncertain grounds. It has not, however, so far as I am informed, been traced any farther back than the Upper Cambrian (Calciferous), and no structural links are known to connect it with either Eozoon or Archæozoon. For this reason it was omitted in the text; but I think it well to mention it here, and to direct attention to it as possibly one of the complex Protozoa which may be traced far back toward the beginnings of life.[57]

Another primitive and generalized genus mentioned in the text is Archæocyathus of Billings, whose headquarters seem to be in the Lower Cambrian, and which may probably be traced farther back.

Mr. Billings described the genus in his "Report on Canadian Fossils" (1861-64), taking A. profundus, from the Lower Cambrian of L'Anse à Loup, on the Labrador coast, in the first instance, as the type.

A few years later, my attention was attracted to this species by specimens presented to me by Mr. Carpenter, a missionary on the Labrador coast, and which Mr. Billings kindly permitted me to compare with his specimens in the Museum of the Geological Survey, collected by the late Mr. Richardson, at L'Anse à Loup, in Labrador, in what were then called Lower Potsdam rocks. Slices of the specimens were made for the microscope, when it appeared that, though they had the general aspect of turbinate corals, like Petraia, etc., they were quite dissimilar in structure, more especially in their porous outer and inner walls and septa (see Fig. 5, p. 35). Yet they could scarcely be referred to the group of porous corals known in much later formations and in the modern seas. Nor could they be referred with much probability to Sponges, as they were composed of solid calcareous plates, which, as was evident from their textures, could not have been originally spicular. One seemed thus shut up to the conclusion that their nearest alliance was with Foraminifera, and if so, they were very large and complex forms of that group, consisting of perforated chambers arranged around a central cavity. I accordingly mentioned them in this connection in 1875, not as closely related to Eozoon, but as apparently showing the existence of very complex foraminiferal forms in the Lower Cambrian.

The specimens thus noticed were altogether calcareous, and were of the species named A. profundus by Mr. Billings. He had, however, referred to the same genus silicified specimens from a later formation, the Calciferous (Upper Cambrian) at Mingan, under the name A. Minganensis, which were subsequently found to be associated with spicules resembling those of lithistid sponges, and which proved to be very different from the Lower Cambrian form, and are now referred to a different genus. The subject had thus become involved in some confusion, and was left in this state by Mr. Billings on his death. I therefore asked my friend, Dr. Hinde, of London, to re-examine my specimens, and at the same time those of the Geological Survey were placed in his hands by Mr. Whiteaves. Hinde also obtained specimens from Lower Cambrian rocks in Sardinia, where they seem to be abundant, and from Spain. He states the results of his examinations very fully in a paper in the Journal of the Geological Society of London.[58] He retains the original name for the older and calcareous form from L'Anse à Loup, separating from it, however, another form, A. Atlanticus of Billings's, which is destitute of distinct radiating septa and acervuline, like the lower part of A. profundus. This he names Spirocyathus. The Mingan species he places with Sponges under the generic name, Archæoscyphia. In this Walcott substantially agrees with Hinde in his "Memoir on the Lower Cambrian Fauna." Both seem to refer Archæocyathus to corals, though admitting its very exceptional and anomalous structure. I think, however, we may still be allowed to entertain some doubts as to the reference to corals, more especially as the skeleton does not seem to have consisted of aragonite, but of ordinary calcite, like that of the Foraminifera. It is in any case a primitive form which seems to be dying out in the Lower Cambrian, and we may hope that it may be traced into the pre-Cambrian, and may form a link connecting the Palæozoic with the Eozoic faunas. In my description of it in "The Dawn of Life" in 1875, I used the following terms:—"To understand Archæocyathus, let us imagine an inverted cone of carbonate of lime from an inch or two to a foot in length, with its point planted in the mud in the bottom of the sea, while its open cup extends upward into the clear water. The lower part buried in the bottom is composed of an irregular network of thick calcareous plates, enclosing chambers communicating with one another. Above this, where the cup expands, its walls are made up of inner and outer plates, perforated with numerous round pores in vertical rows, and connected with each other by vertical partitions also perforated, so as to establish a free communication of the enclosed radiating chambers with each other, as well as with the water within and without. Such a structure might no doubt serve as a skeleton for a coral of somewhat peculiar internal structure, but it might just as well accommodate a protozoan with chambers for its sarcode, and pores for emission of pseudopods, both outwardly and by means of the interior cup, which in that case would represent a funnel like that of Carpenteria, or one of the tubes of Eozoon."

On the whole, when we consider the magnitude and synthetic character of such forms as Cryptozoon, Receptaculites, and Archæocyathus, and their association with generalized types of Crustaceans and Brachiopods, we can scarcely fail to perceive that at the base of the Palæozoic we are leaving the reign of the higher marine invertebrates, and entering on a domain where lower and probably Protozoan forms must be dominant, and so are getting at least within calculable distance of the beginnings of life.

F. Pre-Geological Evolution.

G. Controversies Respecting Eozoon.

In the text (Chapter IX.) I have referred in a cursory manner to these, but have felt that it would be unprofitable to fight the old battles over again, except in so far as the objections raised have suggested new lines of study and investigation. The old objections of Messrs. Rowney, King and Carter were conclusively replied to by the late Dr. Carpenter. The later criticisms of Möbius in his elaborated memoir in "Palæontographica" were in appearance more formidable; but he had evidently entered on the question with imperfect material, and a very defective conception of its extent and meaning. His treatment of it was also marked by unfairness to those who had previously worked at the subject, and by that narrow specialism and captious spirit for which German naturalists are too deservedly celebrated. The difficulties he raised were met at the time, more especially in articles by the present writer in the American Journal of Science, and in the Canadian Naturalist. Möbius, I have no doubt, did his best from his special and limited point of view; but it was a crime which science should not readily pardon or forget, on the part of editors of the German periodical, to publish and illustrate as scientific material a paper which was so very far from being either fair or adequate.

The later objections of Gregory and Lavis are open to similar criticism as imperfect and partial, and as confounding Eozoon with mineral structures which previous writers had carefully distinguished from it. I have stated these points in letters to Nature and to the Council of the Dublin Academy, and have also re-stated the evidence bearing on the animal nature of Eozoon in a series of papers in the Geological Magazine for 1895. I may add here, as apposite to the present condition of the matter, a few remarks referring to the appearance of Eozoon in Dr. Dallinger's new edition of Carpenter's great work on the Microscope,[59] and more especially to his retaining unchanged the description of Eozoon Canadense, as a monument of an important research up to a certain date, while adding a note with reference to the later criticisms of Mr. Gregory.

Dr. Carpenter devoted much time to the study of Eozoon, and brought to bear on it his great experience of foraminiferal forms, and his wonderful powers of manipulating and unravelling difficult structures. After having spent years in studying microscopic slices of Eozoon and the limestones in which it occurs, I have ever felt new astonishment when I saw the manner in which, by various processes of slicing and etching, and by dexterous management of light, he could bring out the structure of specimens often very imperfect. Not long before Dr. Carpenter's death, I had an opportunity to appreciate this in spending a few days with him in studying his more recently acquired specimens, some of them from my own collections, and discussing the new points which they exhibited, and which unhappily he did not live to publish. Some of these new facts, in so far as they related to specimens in our cabinet here, have since that time been noticed in my résumé of the question in the "Memoirs of the Peter Redpath Museum," 1888.

Those who know Dr. Carpenter's powers of investigation will not be astonished that later observers, without his previous preparation and rare insight, and often with only few and imperfect specimens, should have failed to appreciate his results. One is rather surprised that some of them have ventured to state with so great confidence their own negative conclusions in a matter of so much difficulty, and requiring so much knowledge of organic structures in various states of mineralization. For myself, after working fifty years at the microscopic examination of fossils and organic rocks, I feel more strongly than ever the uncertainties and liabilities to error which beset such inquiries.

As an illustration in the case of Eozoon: since the publication of my memoir of 1888, which I had intended to be final and exhaustive as to the main points in so far as I am concerned, I have had occasion to have prepared and to examine about 200 slices of Eozoon from new material; and while most of these have either failed to show the minute structures or have presented nothing new, a few have exhibited certain parts in altogether unexpected perfection, and have shown a prevalence of injection of the canal system by dolomite not previously suspected. I have also observed that unsuitable modes of preparation, notably some of those employed in the preparation of ordinary petrological slices, may fail to disclose organic structures in crystalline limestones when actually present. Since that publication also, the discoveries of Mr. Matthew in the Laurentian of New Brunswick, and the further study of the singular Cambrian forms of the type of Cryptozoon, have opened up new fields of inquiry.

I think it proper to state, in reference to Dr. Dallinger's footnote on the recent paper of Mr. Gregory, that it must not be inferred from it that Mr. Gregory had access to my specimens from Madoc and Tudor, though he no doubt had excellent material from the collections of the Canadian Geological Survey. It might also be inferred from this note that I have regarded the Madoc and Tudor specimens as "Lower Laurentian." The fact is, that I was originally induced in 1865, by the belief of Sir W. E. Logan at that time that these rocks were representatives in a less altered state of the middle part of the Laurentian, to spend some time at Madoc and its vicinity in searching for fossils, but discovered only worm-burrows, spicules, and fragments of Eozoon, which were noticed in the Journal of the Geological Society for 1866. (The more complete specimen from Tudor was found by Vennor in 1866.) On that occasion I satisfied myself fully that the beds are much older than the Cambro-Silurian strata resting on them, unconformably; but I felt disposed to regard them as more probably of the age of some parts of the Huronian of Georgian Bay, which I had explored with a similar purpose under Logan's guidance in 1856.

[In my subsequent notice of the Tudor specimens in "The Dawn of Life," in 1875, I referred to their age as "Upper Laurentian or Huronian"; and I may add, that while it is certain that the beds containing them are pre-Palæozoic, their place in the Eozoic period is still not precisely determined. Work is, however, now in progress which it is hoped may finally settle the age of the "Hastings group" and the old rocks associated with it. I may add that the specimen of Cryptozoon discovered by Mr. Chambers, and of which a portion is represented in the Frontispiece, seems to me to throw a new light on the Tudor specimen. It shows in any case the survival of Cryptozoa similar in form and general appearance to that specimen, as late as the Cambro-Silurian or Ordovician.]

H. Notes to Appendix, December, 1896.