Modiolopsis, Ctenodonta, Orthonota, Palæarca, Lyrodesma, Ambonychia, and Cleidophorus. The Univalves (Gasteropoda) are also very numerous, the two most important genera being Murchisonia (fig. 52) and Pleurotomaria. In both these groups the outer lip of the shell is notched; but the shell in the former is elongated and turreted, whilst in the latter it is depressed. The curious oceanic Univalves known as the Heteropods are also very abundant, the principal forms belonging to Bellerophon and Maclurea. In the former (fig. 53) there is a symmetrical convoluted shell, like that of the Pearly Nautilus in shape, but without any internal partitions, and having the aperture often expanded and notched behind. The species of Maclurea (fig. 54) are found both in North America and in Scotland, and are exclusively confined to the Lower Silurian period, so far as known. They have the shell coiled into a flat spiral, the mouth being furnished with a very curious, thick, and solid lid or " lum." The Lower Silurian Pteropods, or "Winged Snails," are numerous, and belong principally to the genera Theca, Conularia, and Tentaculites, the last-mentioned of these often being extremely abundant in certain strata.

opercu

Lime

stone, America.

Lastly, the Lower Silurian Rocks have yielded a vast number

a

100

Fig. 53-Different views of Bellerophon Argo, Trenton Limestone, Canada.
(After Billings.)

of chambered shells, referable to animals which belong to the same great division as the Cuttle-fishes (the Cephalopoda), and of which the Pearly Nautilus is the only living representative at the present day. In this group of Cephalopods the animal possesses a well-developed external shell, which is divided into chambers by shelly partitions ("septa"). The animal lives in the last-formed and largest chamber of the shell, to

which it is organically connected by muscular attachments. The head is furnished with long muscular processes or "arms,"

Fig. 54.-Different views of Maclurea crenulata, Quebec Group, Newfoundland. (After Billings.)

and can be protruded from the mouth of the shell at will, or again withdrawn within it. We learn, also, from the Pearly Nautilus, that these animals must have possessed two pairs of breathing organs or "gills;" hence all these forms are grouped together under the name of the "Tetrabranchiate" Cephalopods (Gr. tetra, four; bragchia, gill). On the other hand, the ordinary Cuttle-fishes and Calamaries either possess an internal skeleton, or if they have an external shell, it is not chambered; their "arms" are furnished with powerful organs of adhesion in the form of suckers; and they possess only a single pair of gills. For this last reason they are termed the " Dibranchiate" Cephalopods (Gr. dis, twice; bragchia, gill). No trace of the true Cuttle-fishes has yet been found in Lower Silurian deposits; but the Tetrabranchiate group is represented by a great number of forms, sometimes of great size. The principal Lower Silurian genus is the well-known and widely-distributed Orthoceras (fig. 55). The shell in this genus agrees with that of the existing Pearly Nautilus, in consisting of numerous chambers separated by shelly partitions (or septa), the latter being perforated by a tube which runs the whole length of the shell after the last chamber, and is known as the "siphuncle" (fig. 56, s). The last chamber formed is the largest, and in it the animal lives. The chambers behind this are apparently filled with some gas secreted by the animal itself; and these are supposed to act as a kind of float, enabling the creature to move with ease under the weight of its shell. The various airchambers, though the siphuncle passes through them, have no direct connection with one another; and it is believed that the animal has the power of slightly altering its specific gravity, and thus of rising or sinking in the water by driving additional fluid into the siphuncle or partially emptying it. The Ortho

ceras further agrees with the Pearly Nautilus in the fact that the partitions or septa separating the different air-chambers are

simple and smooth, concave in front and convex behind, and devoid of the elaborate lobation which they exhibit in the Ammonites; whilst the siphuncle pierces the septa either in the centre or near it. In the Nautilus, however, the shell is coiled into a flat spiral; whereas in Orthoceras the shell is a straight, longer or shorter cone, tapering behind, and gradually expanding towards its mouth in front. The chief objections to the belief that the animal of the Orthoceras was essentially like that of the Pearly Nautilus are-the comparatively small size of the body-chamber, the often contracted aperture of the mouth, and the enormous size of some specimens of

*This illustration is taken from a rough sketch made by the author many years ago, but he is unable to say from what original source it was copied.

the shell. Thus, some Orthocerata have been discovered measuring ten or twelve feet in length, with a diameter of a foot at the larger extremity. These colossal dimensions certainly make it difficult to imagine that the comparatively small body-chamber could have held an animal large enough to move a load so ponderous as its own shell. To some, this difficulty has appeared so great that they prefer to believe that the Orthoceras did not live in its shell at all, but that its shell was an internal skeleton similar to what we shall find to exist in many of the true Cuttle-fishes. There is something to be said in favour of this view, but it would compel us to believe in the existence in Lower Silurian times of Cuttle-fishes fully equal in size to the giant "Kraken" of fable. It need only be added in this connection that the Lower Silurian rocks have yielded the remains of many other Tetrabranchiate Cephalopods besides Orthoceras. Some of these belong to Cyrtoceras, which only differs from Orthoceras in the bow-shaped form of the shell; others belong to Phragmoceras, Lituites, &c.; and, lastly, we have true Nautili, with their spiral shells, closely resembling the existing Pearly Nautilus.

Whilst all the sub-kingdoms of the Invertebrate animals are represented in the Lower Silurian rocks, no traces of Vertebrate animals have ever been discovered in these ancient deposits, unless the so-called "Conodonts" found by Pander in vast numbers in strata of this age* in Russia should prove to be really of this nature. These problematical bodies are of microscopic size, and have the form of minute, conical, toothshaped spines, with sharp edges, and hollow at the base. Their original discoverer regarded them as the horny teeth of fishes allied to the Lampreys; but Owen came to the conclusion that they probably belonged to Invertebrates. The recent investigation of a vast number of similar but slightly larger bodies, of very various forms, in the Carboniferous rocks of Ohio, has led Professor Newberry to the conclusion that these singular fossils really are, as Pander thought, the teeth of Cyclostomatous fishes. The whole of this difficult question has thus been reopened, and we may yet have to record the first advent of Vertebrate animals in the Lower Silurian.

*

Should

According to Pander, the "Conodonts" are found not only in the Lower Silurian beds, but also in the "Ungulite Grit " (Upper Cambrian), as well as in the Devonian and Carboniferous deposits of Russia. the Conodonts prove to be truly the remains of fishes, we should thus have to transfer the first appearance of Vertebrates to, at any rate, as early a period as the Upper Cambrian.

CHAPTER X.

THE UPPER SILURIAN PERIOD.

Having now treated of the Lower Silurian period at considerable length, it will not be necessary to discuss the succeeding group of the Upper Silurian in the same detail-the more so, as with a general change of species the Upper Silurian animals belong for the most part to the same great types as those which distinguish the Lower Silurian. As compared, also, as regards the total bulk of strata concerned, the thickness of the Upper Silurian is generally very much below that of the Lower Silurian, indicating that they represent a proportionately shorter period of time. In considering the general succession of the Upper Silurian beds, we shall, as before, select Wales and America as being two regions where these deposits are typically developed.

In Wales and its borders the general succession of the Upper Silurian rocks may be taken to be as follows, in ascending order (fig. 57)

(1) The base of the Upper Silurian series is constituted by a series of arenaceous beds, to which the name of "May Hill Sandstone" was applied by Sedgwick. These are succeeded by a series of greenish-grey or pale-grey slates ("Tarannon Shales"), sometimes of great thickness; and these two groups of beds together form what may be termed the "May Hill Group" (Upper Llandovery of Murchison). Though not very extensively developed in Britain, this zone is one very well marked by its fossils; and it corresponds with the "Clinton Group" of North America, in which similar fossils occur. South Wales this group is clearly unconformable to the highest member of the subjacent Lower Silurian (the Llandovery group); and there is reason to believe that a similar, though less conspicuous, physical break occurs very generally between the base of the Upper and the summit of the Lower Silurian.

In

(2) The Wenlock Group succeeds the May Hill group, and constitutes the middle member of the Upper Silurian. At its base it may have an irregular limestone ("Woolhope Limestone"), and its summit may be formed by a similar but thicker calcareous deposit ("Wenlock Limestone"); but the bulk of the group is made up of the argillaceous and shaly strata known as the "Wenlock Shale." In North Wales the Wenlock group is represented by a great accumulation of flaggy and gritty strata (the "Denbighshire Flags and Grits"), and similar beds (the