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THE ANCIENT
LIFE-HISTORY
OF THE EARTH

Chapter 8:

THE CAMBRIAN PERIOD.

The traces of life in the Laurentian period, as we have seen, are but scanty; but the Cambrian Rocks—so called from their occurrence in North Wales and its borders ("Cambria ")—have yielded numerous remains of animals and some dubious plants. The Cambrian deposits have thus a special interest as being the oldest rocks in which occur any number of well-preserved and unquestionable organisms. We have here the remains of the first fauna, or assemblage of animals, of which we have at present knowledge. As regards their geographical distribution, the Cambrian Rocks have been recognised in many parts of the world, but there is some question as to the precise limits of the formation, and we may consider that their most typical area is in South Wales, where they have been carefully worked out, chiefly by Dr Henry Hicks. In this region, in the neighbourhood of the promontory of St David's, the Cambrian Rocks are largely developed, resting upon an ancient ridge of Pre-Cambrian (Laurentian?) strata, and overlaid by the lowest beds of the Lower Silurian. The subjoined sketch-section (fig. 27) exhibits in a general manner the succession of strata in this locality.

From this section it will be seen that the Cambrian Rocks in Wales are divided in the first place into a lower and an upper group. The Lower Cambrian is constituted at the base by a great series of grits, sandstones, conglomerates, and slates, which are known as the "Longmynd group," from their vast development in the Longmynd Hills in Shropshire, and which attain in North Wales a thickness of 8000 feet or more. The Longmynd beds are succeeded by the so-called "Menevian group," a series of sandstones, flags, and grits, about 600 feet in thickness, and containing a considerable number of fossils. The Upper Cambrian series consists in its lower portion of nearly 5000 feet of strata, principally shaly and slaty, which are known as the "Lingula Flags," from the great abundance in them of a shell referable to the genus Lingula. These are followed by 1000 feet of dark shales and flaggy sandstones, which are known as the "Tremadoc slates," from their occurrence near Tremadoc in North Wales; and these in turn are surmounted, apparently quite conformably, by the basement beds of the Lower Silurian.

Page 78 GENERALIZED SECTION OF THE CAMBRIAN ROCKS IN WALES
Fig. 27.
The above may be regarded as giving a typical series of the Cambrian Rocks in a typical locality; but strata of Cambrian age are known in many other regions, of which it is only possible here to allude to a few of the most important. In Scandinavia occurs a well-developed series of Cambrian deposits, representing both the lower and upper parts of the Page 79 formation. In Bohemia, the Upper Cambrian, in particular, is largely developed, and constitutes the so-called "Primordial zone" of Barrande. Lastly, in North America, whilst the Lower Cambrian is only imperfectly developed, or is represented by the Huronian, the Upper Cambrian formation has a wide extension, containing fossils similar in character to the analogous strata in Europe, and known as the "Potsdam Sandstone." The subjoined table shows the chief areas where Cambrian Rocks are developed, and their general equivalency:

Like all the older Palæozoic deposits, the Cambrian Rocks, though by no means necessarily what would be called actually "metamorphic," have been highly cleaved, and otherwise altered from their original condition. Owing partly to their indurated state, and partly to their great antiquity, they are usually found in the heart of mountainous districts, which have undergone great disturbance, and have been subjected to an enormous amount of denudation. In some cases, as in the Longmynd Hills in Shropshire, they form low rounded elevations, largely covered by pasture, and with few or no elements of sublimity. In other cases, however, they rise into bold and rugged mountains, girded by precipitous cliffs. Industrially, the Cambrian Rocks are of interest, if only for the reason that the celebrated Welsh slates of Llanberis are derived from highly-cleaved beds of this age. Taken as a whole, the Cambrian formation is essentially composed of arenaceous and Page 80 muddy sediments, the latter being sometimes red, but more commonly nearly black in colour. It has often been supposed that the Cambrians are a deep-sea deposit, and that we may thus account for the few fossils contained in them; but the paucity of fossils is to a large extent imaginary, and some of the Lower Cambrian beds of the Longmynd Hills would appear to have been laid down in shallow water; as they exhibit rain-prints, sun-cracks, and ripple-marks—incontrovertible evidence of their having been a shore-deposit. The occurrence, of innumerable worm-tracks and burrows in many Cambrian strata is also a proof of shallow-water conditions; and the general absence of limestones, coupled with the coarse mechanical nature of many of the sediments of the Lower Cambrian, maybe taken as pointing in the same direction.

The life of the Cambrian, though not so rich as in the succeeding Silurian period, nevertheless consists of representatives of most of the great classes of invertebrate animals. The coarse sandy deposits of the formation, which abound more particularly towards its lower part, naturally are to a large extent barren of fossils; but the muddy sediments, when not too highly cleaved, and especially towards the summit of the group, are replete with organic remains. This is also the case, in many localities at any rate, with the finer beds of the Potsdam Sandstone in America. Limestones are known to occur in only a few areas (chiefly in America), and this may account for the apparent total absence of corals. It is, however, interesting to note that, with this exception, almost all the other leading groups of Invertebrates are known to have come into existence during the Cambrian period.

Of the land-surfaces of the Cambrian period we know nothing; and there is, therefore, nothing surprising in the fact that our acquaintance with the Cambrian vegetation is confined to some marine plants or sea-weeds, often of a very obscure and problematical nature. The "Fucoidal Sandstone" of Sweden, and the "Potsdam Sandstone" of North America, have both yielded numerous remains which have been regarded as markings left by sea-weeds or "Fucoids;" but these are highly enigmatical in their characters, and would, in many instances, seem to be rather referable to the tracks and burrows of marine worms. The first-mentioned of these formations has also yielded the curious, furrowed and striated stems which have been described as a kind of land-plant under the name of Eopkyton (fig. 28). It cannot be said, however, that the vegetable origin of these singular bodies has been satisfactorily proved. Lastly, there are found in certain green Page 81 and purple beds of Lower Cambrian age at Bray Head, Wicklow, Ireland, some very remarkable fossils, which are well known under
Fig. 28.—Fragment of Eophyton Linneanum, a supposed land-plant. Lower Cambrian, Sweden, of the natural size. the name of Oldhamia, but the true nature of which is very doubtful. The commonest form of Oldhamia (fig. 29) consists of a thread-like stem or axis, from which spring at regular intervals bundles of short filamentous branches in a fan-like manner. In the locality where it occurs, the fronds of Oldhamia are very abundant, and are spread over the surfaces of the strata in tangled layers. That it is organic is certain, and that it is a calcareous sea-weed is probable; but it may possibly belong to the sea-mosses (Polyzoa), or to the sea-firs (Sertularians).

Amongst the lower forms of animal life (Protozoa), we find the Sponges represented by the curious bodies, composed of netted fibres, to which the name of Protospongia has been given (fig. 32, a); and the comparatively gigantic, conical, or Page 82 cylindrical fossils termed Archœocyathus by Mr Billings are certainly referable either to the Foraminifera
Fig. 29.—A portion of Oldhamia antiqua, Lower Cambrian, Wicklow, Ireland, of the natural size. (After Salter.) or to the Sponges. The almost total absence of limestones in the formation may be regarded as a sufficient explanation of the fact that the Foraminifera are not more largely and unequivocally represented; though the existence of greensands in the Cambrian beds of Wisconsin and Tennessee may be taken as an indication that this class of animals was by no means wholly wanting. The same fact may explain the total absence of corals, so far as at present known.

The group of the Echinodermata (Sea-lilies, Sea-urchins, and their allies) is represented by a few forms, which are principally of interest as being the earliest-known examples of the class. It is also worthy of note that these precursors of a group which subsequently attains such geological importance, are referable to no less than three distinct orders—the Crinoids or Sea-lilies, represented by a species of Dendrocrinus; the Cystideans by Protocystites; and the Star-fishes by Palasterina and some other forms. Only the last of these groups, however, appears to occur in the Lower Cambrian.

The Ringed-worms (Annelida), if rightly credited with all the remains usually referred to them, appear to have swarmed in the Cambrian seas. Being soft-bodied, we do not find the actual worms themselves in the fossil condition, but we have, nevertheless, abundant traces of their existence. In some cases we find vertical burrows of greater or less depth, often expanded towards their apertures, in which the worm must have actually lived (fig. 30), as various species do at the present day. In these cases, the tube must have been rendered more or less permanent by receiving a coating of mucus, or perhaps a genuine membranous secretion, from the body of the animal; and it may be found quite empty, or occupied by a cast of sand or mud. Of this nature are the burrows which have been described under the names of Scolithus and Scolecoderma, and probably the Histioderma of the Lower Cambrian Page 83 of Ireland. In other cases, as in Arenicolites (fig. 32, b), the worm seems to have inhabited a double
Fig. 30.—Annelide-burrows (Scolithus linearus) from the Potsdam Sandstone of Canada, of the natural size. (After Billings.) burrow, shaped like the letter U, and having two openings placed close together on the surface of the stratum. Thousands of these twin-burrows occur in some of the strata of the Longmynd, and it is supposed that the worm used one opening to the burrow as an aperture of entrance, and the other as one of exit. In other cases, again, we find simply the meandering trails caused by the worm dragging its body over the surface of the mud. Markings of this kind are commoner in the Silurian Rocks, and it is generally more or less doubtful whether they may not have been caused by other marine animals, such as shellfish, whilst some of them have certainly nothing whatever to do with the worms. Lastly, the Cambrian beds often show twining cylindrical bodies, commonly more or less matted together, and not confined to the surfaces of the strata, but passing through them. These have often been regarded as the remains of sea-weeds, but it is more probable that they represent casts of the underground burrows of worms of similar habits to the common lob-worm (Arenicola) of the present day.

The Articulate animals are numerously represented in the Cambrian deposits, but exclusively by the class of Crustaceans. Some of these are little double-shelled creatures, resembling our living water-fleas (Ostracoda). A few are larger forms, and belong to the same group as the existing brine-shrimps and fairy-shrimps (Phyllopoda). One of the most characteristic Page 84 of these is the Hymenocaris vermicauda of the Lingula Flags (fig. 32, d). By far the larger number of the Cambrian Crustacea belong, however, to the remarkable and wholly extinct group of the Trilobites. These extraordinary animals must have literally swarmed in the seas of the later portion of this and the whole of the succeeding period; and they survived in greatly diminished numbers till the earlier portion of the Carboniferous period. They died out, however, wholly before the close of the Palæozoic epoch, and we have no Crustaceans at the present day which can be considered as their direct representatives. They have, however, relationships of a more or less intimate character with the existing groups of the Phyllopods, the King-crabs (Limulus), and the Isopods ("Slaters," Wood-lice, &c.) Indeed, one member of the last-mentioned order, namely, the Serolis of the coasts of Patagonia, has been regarded as the nearest living ally of the Trilobites. Be this as it may, the Trilobites possessed a skeleton which, though capable of undergoing almost endless variations, was wonderfully constant in its pattern of structure, and we may briefly describe here the chief features of this.

The upper surface of the body of a Trilobite was defended by a strong shell or "crust," partly horny and partly calcareous in its composition. This shell (fig. 31) generally exhibits a very distinct "trilobation" or division into three longitudinal lobes, one central and two lateral. It also exhibits a more important and more fundamental division into three transverse portions, which are so loosely connected with one another as very commonly to be found separate. The first and most anterior of these divisions is a shield or buckler which covers the head; the second or middle portion is composed of movable rings covering the trunk ("thorax "); and the third is a shield which covers the tailor "abdomen." The head-shield (fig. 31, e) is generally more or less semicircular in shape; and its central portion, covering the stomach of the animal, is usually strongly elevated, and generally marked by lateral furrows. A little on each side of the head are placed the eyes, which are generally crescentic in shape, and resemble the eyes of insects and many existing Crustaceans in being "compound," or made up of numerous simple eyes aggregated together. So excellent is the state of preservation of many specimens of Trilobites, that the numerous individual lenses of the eyes have been uninjured, and as many as four hundred have been counted in each eye of some forms. The eyes may be supported upon prominences, but they are never carried on movable stalks (as they are in the existing lobsters and crabs); and Page 85 in some of the Cambrian Trilobites, such as the little Agnosti (fig. 31 g), the animal was blind. The lateral portions of the head-shield are usually separated from the central portion by a peculiar
Fig. 31.—Cambrian Trilobites: a, Paradoxides Bohemicus, reduced in size; b, Ellipsocephalus Hoffi; c, Sao hirsuta; d, Conocorypke Sultzeri (all the above, together with fig. g, are from the Upper Cambrian or "Primordial Zone" of Bohemia); e, Head-shield of Dikellocephalus Celticus, from the Lingula Flags of Wales; f, Head-shield of Conocoryphe Matthewi, from the Upper Cambrian (Acadian Group) of New Brunswick; g, Agnostus rex, Bohemia; h, Tail-shield of Dikellocephalus Minnesotensis, from the Upper Cambrian (Potsdam Sandstone) of Minnesota. (After Barrande, Dawson, Salter, and Dale Owen.) line of division (the so-called "facial suture") on each side; but this is also wanting in some of the Cambrian species. The backward angles of the head-shield, also, are often prolonged into spines, which sometimes reach a great length. Following the head-shield behind, we have a portion of the body which is composed of movable segments or "body-rings," and which is technically called the "thorax," Ordinarily, this region is strongly trilobed, and each ring consists of a central convex portion, and of two flatter side-lobes. The number of body-rings in the thorax is very variable (from two to twenty-six), but is fixed for the adult forms of each group of the Trilobites. The young forms have much fewer rings than the full-grown ones; and it is curious to find that the Cambrian Page 86 Trilobites very commonly have either a great many rings (as in Paradoxides, fig. 31, a), or else very few (as in Agnostus, fig. 31, g). In some instances, the body-rings do not seem to have been so constructed as to allow of much movement, but in other cases this region of the body is so flexible that the animal possessed the power of rolling itself up completely, like a hedgehog; and many individuals have been permanently preserved as fossils in this defensive condition. Finally, the body of the Trilobite was completed by a tail-shield (technically termed the "pygidium"), which varies much in size and form, and is composed of a greater or less number of rings, similar to those which form the thorax, but immovably amalgamated with one another (fig. 31, h).

The under surface of the body in the Trilobites appears to have been more or less entirely destitute of hard structures, with the exception of a well-developed upper lip, in the form of a plate attached to the inferior side of the head-shield in front. There is no reason to doubt that the animal possessed legs; but these structures seem to have resembled those of many living Crustaceans in being quite soft and membranous. This, at any rate, seems to have been generally the case; though structures which have been regarded as legs have been detected on the under surface of one of the larger species of Trilobites. There is also, at present, no direct evidence that the Trilobites possessed the two pairs of jointed feelers ("antennæ") which are so characteristic of recent Crustaceans.

The Trilobites vary much in size, and the Cambrian formation presents examples of both the largest and the smallest members of the order. Some of the young forms may be little bigger than a millet-seed, and some adult examples of the smaller species (such as Agnostus) may be only a few lines in length; whilst such giants of the order as Paradoxides and Asaphus may reach a length of from one to two feet. Judging from what we actually know as to the structure of the Trilobites, and also from analogous recent forms, it would seem that these ancient Crustaceans were mud-haunting creatures, denizens of shallow seas, and affecting the soft silt of the bottom rather than the clear water above. Whenever muddy sediments are found in the Cambrian and Silurian formations, there we are tolerably sure to find Trilobites, though they are by no means absolutely wanting in limestones. They appear to have crawled out upon the sea-bottom, or burrowed in the yielding mud, with the soft under surface directed downwards; and it is probable that they really derived their nutriment from the organic matter contained in the ooze amongst which they Page 87 lived. The vital organs seem to have occupied the central lobe of the skeleton, by which they were protected; and a series of delicate leaf-like paddles, which probably served as respiratory organs, would appear to have been carried on the under surface of the thorax. That they had their enemies may be regarded as certain; but we have no evidence that they were furnished with any offensive weapons, or, indeed, with any means of defence beyond their hard crust, and the power, possessed by so many of them, of rolling themselves into a ball. An additional proof of the fact that they for the most part crawled along the sea-bottom is found in the occurrence of tracks and markings of various kinds, which can hardly be ascribed to any other creatures with any show of probability. That this is the true nature of some of the markings in question cannot be doubted at all; and in other cases no explanation so probable has yet been suggested. If, however, the tracks which have been described from the Potsdam Sandstone of North America under the name of Protichnites are really due to the peregrinations of some Trilobite, they must have been produced by one of the largest examples of the order.

As already said, the Cambrian Rocks are very rich in the remains of Trilobites. In the lowest beds of the series (Longmynd Rocks), representatives of some half-dozen genera have now been detected, including the dwarf Agnostus and the giant Paradoxides. In the higher beds, the number both of genera and species is largely increased; and from the great comparative abundance of individuals, the Trilobites have every right to be considered as the most characteristic fossils of the Cambrian period,—the more so as the Cambrian species belong to peculiar types, which, for the most part, died out before the commencement of the Silurian epoch.

All the remaining Cambrian fossils which demand any notice here are members of one or other division of the great class of the Mollusca, or "Shell-fish" properly so called. In the Lower Cambrian Rocks the Lamp-shells (Brachiopoda) are the principal or sole representatives of the class, and appear chiefly in three interesting and important types—namely, Lingulella, Discina, and Obolella. Of these the last (fig. 32, i) is highly characteristic of these ancient deposits; whilst Discina is one of those remarkable persistent types which, commencing at this early period, has continued to be represented by varying forms through all the intervening geological formations up to the present day. Lingulella (fig. 32, c), again, is closely allied to the existing "Goose-bill" Lamp-shell (Lingula anatina), and thus presents us with another example of an extremely long-lived Page 88 type. The Lingulellœ and their successors; the Lingulœ, are singular in possessing a shell which is of a horny texture, and contains but a small proportion of calcareous matter. In the Upper Cambrian Rocks, the Lingulellœ become much more abundant, the broad satchel-shaped species known as L. Davisii (fig. 32, e) being so abundant that one of the great divisions of the Cambrian is termed the "Lingula Flags." Here, also, we meet for the first time with examples of the genus Orthis (fig. 32, f, k, l) a characteristic Palæozoic type of the Brachiopods, which is destined to undergo a vast extension in later ages.

Fig. 32.—Cambrian Fossils: a, Protospongia fenestrata, Menevian Group; b, Arenicolites didymus, Longmynd Group; c, Lingulella ferruginea, Longmynd and Menevian, enlarged; d, Hymenocaris vermicauda, Lingula Flags; e, Lingulella Davisii, Lingula Flags; f, Orthis lenticularis, Lingula Flags; g, Theca Davidii, Tremadoc Slates; h, Modiolopsis Solvensis, Tremadoc Slates; i, Obolela sagittalis, interior of valve, Menevian; j, Exterior of the same; k, Orthis Hicksii, Menevian; l, Cast of the same; m, Olenus micrurus, Lingula Flags. (Alter Salter, Hicks, and Davidson.)

Fig. 33.—Fragment of Dictyonema sociale, considerably enlarged, showing the horny branches, with their connecting cross-bars, and with a row of cells on each side. (Original.)

Of the higher groups of the Mollusca the record is as yet but scanty. In the Lower Cambrian, we have but the thin, fragile, dagger-shaped shells of the free-swimming oceanic Molluscs or "Winged-snails" (Pteropoda), of which the most characteristic is the genus Theca (fig. 32, g). In the Upper Cambrian, in addition to these, we have a few Univalves (Gasteropoda), and, thanks to the researches of Dr Hicks, quite a small assemblage of Bivalves (Lamellibranchiata), though these are mostly of no great dimensions (fig. 32, h). Of the chambered Cephalopoda (Cuttle-fishes and their allies), we have but Page 89 few traces; and these wholly confined to the higher beds of the formation. We meet, however, with examples of the wonderful genus Orthoceras, with its straight, partitioned shell, which we shall find in an immense variety of forms in the Silurian rocks. Lastly, it is worthy of note that the lowest of all the groups of the Mollusca—namely, that of the Sea-mats, Sea-mosses, and Lace-corals (Polyzoa)—is only doubtfully known to have any representatives in the Cambrian, though undergoing a large and varied development in the Silurian deposits.

An exception, however, may with much probability be made to this statement in favour of the singular genus Dictyonema (fig. 33), which is highly characteristic of the highest Cambrian beds (Tremadoc Slates). This curious fossil occurs in the form of fan-like or funnel-shaped expansions, composed of slightly-diverging horny branches, which are united in a net-like manner by numerous delicate cross-bars, and exhibit a row of little cups or cells, in which the animals were contained, on each side. Dictyonema has generally been referred to the Graptolites; but it has a much greater affinity with the plant-like Sea-firs (Sertularians) or the Sea-mosses (Polyzoa), and the balance of evidence is perhaps in favour of placing it with the latter.

LITERATURE.

The following are the more important and accessible works and memoirs which may be consulted in studying the stratigraphical and palæontological relations of the Cambrian Rocks:—

In the above list, allusion has necessarily been omitted to numerous works and memoirs on the Cambrian deposits of Sweden and Norway, Central Europe, Russia, Spain, and various parts of North America, as well as to a number of important papers on the British Cambrian strata by various well-known observers. Amongst these latter may be mentioned memoirs by Prof. Phillips, and Messrs Salter, Hicks, Belt, Plant, Homfray, Ash, Holl, &c.