1911 Encyclopædia Britannica/Devonian System
DEVONIAN SYSTEM, in geology, the name applied to series of stratified fossiliferous and igneous rocks that were formed during the Devonian period, that is, in the interval of time between the close of the Silurian period and the beginning of the Carboniferous; it includes the marine Devonian and an estuarine Old Red Sandstone series of strata. The name "Devonian" was introduced in 1829 by Sir R. Murchison and A. Sedgwick to describe the older rocks of Cornwall and Devon which W. Lonsdale had shown, from an examination of the fossils, to be intermediate between the Silurian and Carboniferous. The same two workers also carried on further researches upon the same rocks of the European continent, where already several others, F. Roemer, H. E. Beyrich, &c., were endeavouring to elucidate the succession of strata in this portion of the "Transition Series." The labours of these earlier workers, including in addition to those already mentioned, the brothers F. and G. von Sandberger, A. Dumont, J. Gosselet, E. J. A. d'Archiac, E. P. de Verneuil and H. von Dechen, although somewhat modified by later students, formed the foundation upon which the modern classification of the Devonian rocks is based.
Stratigraphy of the Devonian Facies.
Notwithstanding the fact that it was in Devonshire and Cornwall that the Devonian rocks were first distinguished, it is in central Europe that the succession of strata is most clearly made out, and here, too, their geological position was first indicated by the founders of the system, Sedgwick and Murchison.
Continental Europe.—Devonian rocks occupy a large area in the centre of Europe, extending from the Ardennes through the south of Belgium across Rhenish Prussia to Darmstadt. They are best known from the picturesque gorges which have been cut through them by the Rhine below Bingen and by the Moselle below Treves. They reappear from under younger formations in Brittany, in the Harz and Thuringia, and are exposed in Franconia, Saxony, Silesia, North Moravia and eastern Galicia. The principal subdivisions of the system in the more typical areas are indicated in Table I.
This threefold subdivision, with a central mass of calcareous strata, is traceable westwards through Belgium (where the Calcaire de Givet represents the Stringocephalus limestone of the Eifel) and eastwards into the Harz. The rocks reappear with local petrographical modifications, but with a remarkable persistence of general palaeontological characters, in Eastern Thuringia, Franconia, Saxony, Silesia, the north of Moravia and East Galicia. Devonian rocks have been detected among the crumpled rocks of the Styrian Alps by means of the evidence of abundant corals, cephalopods, gasteropods, lamellibranchs and other organic remains. Perhaps in other tracts of the Alps, as well as in the Carpathian range, similar shales, limestones and dolomites, though as yet unfossiliferous, but containing ores of silver, lead, mercury, zinc, cobalt and other metals, may be referable to the Devonian system.
In the centre of Europe, therefore, the Devonian rocks consist of a vast thickness of dark-grey sandy and shaly rocks, with occasional seams of limestone, and in particular with one thick central calcareous zone. These rocks are characterized in the lower zones by numerous broad-winged spirifers and by peculiar trilobites (Phacops, Homalonotus, &c.) which, though generically like those of the Silurian system, are specifically distinct. The central calcareous zone abounds in corals and crinoids as well as in numerous brachiopods. In the highest bands a profusion of coiled cephalopods (Clymenia) occurs in some of the limestones, while the shales are crowded with a small but characteristic ostracod crustacean (Cypridina). Here and there traces of fishes have been found, more especially in the Eifel, but seldom in such a state of preservation as to warrant their being assigned to any definite place in the zoological scale. Subsequently, however, E. Beyrich has described from Gerolstein in the Eifel an undoubted species of Pterichthys, which, as it cannot be certainly identified with any known form, he names P. Rhenanus. A Coccosteus has been described by F. A. Roemer from the Harz, and still later one has been cited from Bicken near Herborn by V. Koenen; but, as Beyrich points out, there may be some doubt as to whether the latter is not a Pterichthys. A Ctenacanthus, seemingly undistinguishable from the C. Bohemicus of Barrande's Étage G, has also been obtained from the Lower Devonian "Nereitenschichten" of Thuringia. The characteristic Holoptychius nobilissimus has been detected in the Psammite de Condroz, which in Belgium forms a characteristic sandy portion of the Upper Devonian rocks. These are interesting facts, as helping to link the Devonian and Old Red Sandstone types together. But they are as yet too few and unsupported to warrant any large deduction as to the correlations between these types.
It is in the north-east of Europe that the Devonian and Old Red Sandstone appear to be united into one system, where the limestones and marine organisms of the one are interstratified with the fish-bearing sandstones and shales of the other. In Russia, as was shown in the great work Russia and the Ural Mountains by Murchison, De Verneuil and Keyserling, rocks intermediate between the Upper Silurian and Carboniferous Limestone formations cover an extent of surface larger than the British Islands. This wide development arises not from the thickness but from the undisturbed horizontal character of the strata. Like the Silurian formations described elsewhere, they remain to this day nearly as flat and unaltered as they were originally laid down. Judged by mere vertical depth, they present but a meagre representative of the massive Devonian greywacke and limestone of Germany, or of the Old Red Sandstone of Britain. Yet vast though the area is over which they form the surface rock, it is probably only a small portion of their total extent; for they are found turned up from under the newer formations along the flank of the Ural chain. It would thus seem that they spread continuously across the whole breadth of Russia in Europe. Though almost everywhere undisturbed, they afford evidence of some terrestrial oscillation between the time of their formation and that of the Silurian rocks on which they rest, for they are found gradually to overlap Upper and Lower Silurian formations.
|Limestone of Etrœungt.
Psammites of Condroz
Slates of Famenne
Pön sandstone (Sauerland).
Crumbly limestone (Kramen-
zelkalk) with Clymenia.
Neheim slates in Sauerland,
and diabases, tuffs, &c., in
|Slates of Rostellec.||Cypridina slates.
and limestone of
|Slates of Matagne.
Limestones, marls and
shale of Frasne, and
red marble of
|Adorf limestone of Waldeck
and shales with Goniatites
(Eifel and Aix) =
Marls, limestone and dolomite
with Rhynchonella cuboides
(Flinz in part).
Iberg limestone of Dillmulde.
|Limestone of Cop-
Choux and green
slates of Travuliors.
|Iberg limestone and
stone; also Adorf
limestone and shales
|Limestone of Givet.||Stringocephalus limestone,
ironstone of Brilon and
Upper Lenne shales, crinoidal
limestone of Eifel, red
sandstones of Aix.
Tuffs and diabases of Brilon
Red conglomerate of Aix.
|H2 (of Barrande)
with Flaser and
|Calceola slates and
limestones of Couvin.
Greywacke with Spirifer
|Calceola beds, Wissenbach
slates, Lower Lenne beds,
Güntroder limestone and
clay slate of Lahnmulde,
|Slates of Porsguen,
greywacke of Fret.
Nereite slates, slates
of Wieda and lime-
stones of Hasselfeld.
|Coblentzien||Greywacke of Hierges.
Shales and conglomer-
ate of Burnot with
quartzite, of Bierlé
and red slates of
of Vireux, greywacke
of Montigny, sand-
stone of Anor.
|Upper Coblentz slates.
Red sandstone of Eifel,
Coblentz quartzite, lower
Hunsrück and Siegener
greywacke and slates.
Taunus quartzite and
of Erbray, Brulon,
Viré and Néhou,
greywacke of Faou,
|F2 of Barrande.
|Haupt quartzite (of
Lossen) = Rammelsberg
slates, Schallker slates =
Hercynian slates and
|Gédinnien||Slates of St Hubert and
and Fooz, slates of
Mondrepuits, arkose of
|Slates of Gédinne.||Slates and quartzites
The chief interest of the Russian rocks of this age lies in the fact, first signalized by Murchison and his associates, that they unite within themselves the characters of the Devonian and the Old Red Sandstone types. In some districts they consist largely of limestones, in others of red sandstones and marls. In the former they present molluscs and other marine organisms of known Devonian species; in the latter they afford remains of fishes, some of which are specifically identical with those of the Old Red Sandstone of Scotland. The distribution of these two palaeontological types in Russia is traced by Murchison to the lithological characters of the rocks, and consequent original diversities of physical conditions, rather than to differences of age. Indeed cases occur where in the same band of rock Devonian shells and Old Red Sandstone fishes lie commingled. In the belt of the formation which extends southwards from Archangel and the White Sea, the strata consist of sands and marls, and contain only fish remains. Traced through the Baltic provinces, they are found to pass into red and green marls, clays, thin limestones and sandstones, with beds of gypsum. In some of the calcareous bands such fossils occur as Orthis striatula, Spiriferina prisca, Leptaena productoides, Spirifer calcaratus, Spirorbis omphaloides and Orthoceras subfusiforme. In the higher beds Holoptychius and other well-known fishes of the Old Red Sandstone occur. Followed still farther to the south, as far as the watershed between Orel and Voronezh, the Devonian rocks lose their red colour and sandy character, and become thin-bedded yellow limestones, and dolomites with soft green and blue marls. Traces of salt deposits are indicated by occasional saline springs. It is evident that the geographical conditions of the Russian area during the Devonian period must have closely resembled those of the Rhine basin and central England during the Triassic period. The Russian Devonian rocks have been classified in Table II. There is an unquestionable passage of the uppermost Devonian rocks of Russia into the base of the Carboniferous system.
|North-West Russia.||Central Russia.||Petchoraland.||Ural Region.|
Spirifer Verneuili and
|Limestones with Arca
Limestones with Sp.
Verneuili and Sp.
|Domanik slates and
limestones with Sp.
and Rhynchonella cuboides
|Middle.||Dolomites and limestones
|Limestones and slates
with Sp. Anossofi
Limestones and slates with
|Lower sandstone (Old Red).|
|Lower.||Absent.||Limestones and slates of the
Yuresan and Ufa rivers,
slate and quartzite,
marble of Byclaya and
of Bogoslovsk, phyllitic
schists and quartzite.
The Lower Devonian of the Harz contains a fauna which is very different from that of the Rhenish region; to this facies the name "Hercynian" has been applied, and the correlation of the strata has been a source of prolonged discussion among continental geologists. A similar fauna appears in Lower Devonian of Bohemia, in Brittany (limestone of Erbray) and in the Urals. The Upper Devonian of the Harz passes up into the Culm.
In the eastern Thuringian Fichtelgebirge the upper division is represented by Clymenia limestone and Cypridina slates with Adorf limestone, diabase and Planschwitzer tuff in the lower part. The middle division has diabases and tuffs at the top with Tentaculite and Nereite shales and limestones below. The upper part of the Lower Devonian, the sandy shale of Steinach, rests unconformably upon Silurian rocks. In the Carnic Alps are coral reef limestones, the equivalents of the Iberg limestone, which attain an enormous thickness; these are underlain by coral limestones with fossils similar to those of the Konjeprus limestone of Bohemia; below these are shales and nodular limestones with goniatites. The Devonian rocks of Poland are sandy in the lower, and more calcareous in the upper parts. They are of interest because while the upper portions agree closely with the Rhenish facies, from the top of the Coblentzien upwards, in the sandy beds near the base Old Red Sandstone fishes (Coccosteus, &c.) are found. In France Devonian rocks are found well developed in Brittany, as indicated in the table, also in Normandy and Maine; in the Boulonnais district only the middle and upper divisions are known. In south France in the neighbourhood of Cabrières, about Montpellier and in the Montagne Noire, all three divisions are found in a highly calcareous condition. Devonian rocks are recognized, though frequently much metamorphosed, on both the northern and southern flanks of the Pyrenees; while on the Spanish peninsula they are extensively developed. In Asturias they are no less than 3280 ft. thick, all three divisions and most of the central European subdivisions are present. In general, the Lower Devonian fossils of Spain bear a marked resemblance to those of Brittany.
Asia.—From the Ural Mountains eastward, Devonian rocks have been traced from point to point right across Asia. In the Altai Mountains they are represented by limestones of Coblentzien age with a fauna possessing Hercynian features. The same features are observed in the Devonian of the Kougnetsk basin, and in Turkestan. Well-developed quartzites with slates and diabases are found south of Yarkand and Khotan. Middle and Upper Devonian strata are widespread in China. Upper Devonian rocks are recorded from Persia, and from the Hindu Kush on the right bank of the Chitral river.
England.—In England the original Devonian rocks are developed in Devon and Cornwall and west Somerset. In north Devonshire these rocks consist of sandstones, grits and slates, while in south Devon there are, in addition, thick beds of massive limestone, and intercalations of lavas and tuffs. The interpretation of the stratigraphy in this region is a difficult matter, partly on account of the absence of good exposures with fossils, and partly through the disturbed condition of the rocks. The system has been subdivided as shown in Table III.
|North Devon and West
|Upper.||Pilton group. Grits, slates
and thin limestones.
Baggy group. Sandstones
Pickwell Down group.
Dark slates and grits.
Morte slates (?).
Red and green Entomis slates
Red and grey slates with
Chudleigh goniatite limestone
Petherwyn beds (Frasnien).
|Middle.||Ilfracombe slates with
lenticles of limestone.
Combe Martin grits and
|Torquay and Plymouth
limestones and Ashprington
volcanic series. (Givétien
Slates and limestones of
|Lower.||Hangman grits and slates.
Lynton group, grits and
Foreland grits and slates.
|Looe beds (Cornwall).
Meadfoot, Cockington and
Warberry series of slates
and greywackes. (Coblentzien
The fossil evidence clearly shows the close agreement of the Rhenish and south Devonshire areas. In north Devonshire the Devonian rocks pass upward without break into the Culm.
North America.—In North America the Devonian rocks are extensively developed; they have been studied most closely in the New York region, where they are classified according to Table IV.
The classification below is not capable of application over the states generally and further details are required from many of the regions where Devonian rocks have been recognized, but everywhere the broad threefold division seems to obtain. In Maryland the following arrangement has been adopted—(1) Helderberg = Coeymans; (2) Oriskany; (3) Romney = Erian; (4) Jennings = Genesee and Portage; (5) Hampshire = Catskill in part. In the interior the Helderbergian is missing and the system commences with (1) Oriskany, (2) Onondaga, (3) Hamilton, (4) Portage (and Genesee), (5) Chemung.
The Helderbergian series is mainly confined to the eastern part of the continent; there is a northern development in Maine, and in Canada (Gaspé, New Brunswick, Nova Scotia and Montreal); an Appalachian belt, and a lower Mississippian region. The series as a whole is mainly calcareous (2000 ft. in Gaspé), and thins out towards the west. The fauna has Hercynian affinities. The Oriskany formation consists largely of coarse sandstones; it is thin in New York, but in Maryland and Virginia it is several hundred feet thick. It is more widespread than the underlying Helderbergian. The Lower Devonian appears to be thick in northern Maine and in Gaspé, New Brunswick and Nova Scotia, but neither the palaeontology nor the stratigraphy has been completely worked out.
In the Middle Devonian the thin clastic deposits at the base, Esopus and Schoharie grits, have not been differentiated west of the Appalachian region; but the Onondaga limestones are much more extensive. The Erian series is often described as the Hamilton series outside the New York district, where the Marcellus shales are grouped together with the Hamilton shales, and numerous local subdivisions are included, as in Ohio, Kentucky and Tennessee. The rocks are mostly shales or slates, but limestones predominate in the western development. In Pennsylvania the Hamilton series is from 1500 ft. to 5000 ft. thick, but in the more calcareous western extension it is much thinner. The Marcellus shales are bituminous in places.
The Senecan series is composed of shallow-water deposits; the Tully limestone, a local bed in New York, thins out in places into a layer of pyrites which contains a remarkable dwarfed fauna. The bituminous Genesee shales are thickest in Pennsylvania (300 ft.); 25 ft. on Lake Erie. The shales and sandstones of the Portage formation reach 1000 ft. to 1400 ft. in western New York. In the Chautauquan series the Chemung formation is not always clearly separable from the Portage beds, it is a sandstone and conglomerate formation which reaches its maximum thickness (8000 ft.) in Pennsylvania, but thins rapidly towards the west. In the Catskill region the Upper Devonian has an Old Red facies—red shales and sandstones with a freshwater and brackish fauna.
|Upper.||Chautauquan.||Chemung beds with Catskill
as a local facies.
|Senecan.||Portage beds (Naples, Ithaca
and Oneonta shales as local
Esopus grit (Caudagalli grit).
New Scotland beds.
Although the correlation of the strata has only advanced a short distance, there is no doubt as to the presence of undifferentiated Devonian rocks in many parts of the continent. In the Great Plains this system appears to be absent, but it is represented in Colorado, Utah, Nevada, Wyoming, Montana, California and Arizona; Devonian rocks occur between the Sierras and the Rocky Mountains, in the Arbuckle Mountains of Oklahoma and in Texas. In the western interior limestones predominate; 6000 ft. of limestone are found at Eureka, Nevada, beneath 2000 ft. of shale. On the Pacific coast metamorphism of the rocks is common, and lava-flows and tuffs occur in them.
In Canada, besides the occurrences previously mentioned in the eastern region, Devonian strata are found in considerable force along the course of the Mackenzie river and the Canadian Rockies, whence they stretch out into Alaska. It is probable, however, that much that is now classed as Devonian in Canada will prove on fossil evidence to be Carboniferous.
South America, Africa, Australia, &c.—In South America the Devonian is well developed; in Argentina, Bolivia, Brazil, Peru and the Falkland Islands, the palaeontological horizon is about the junction of the Lower and Middle divisions, and the fauna has affinities with the Hamilton shales of North America. Nearly allied to the South American Devonian is that of South Africa, where they are represented by the Bokkeveld beds in the Cape system. In Australia we find Lower Devonian consisting of coarse littoral deposits with volcanic rocks; and a Middle division with coral limestones in Victoria, New South Wales and Queensland; an Upper division has also been observed. In New Zealand the Devonian is well exposed in the Reefton mining field; and it has been suggested that much of the highly metamorphosed rock may belong to this system.
Stratigraphy of the Old Red Sandstone Facies.
The Old Red Sandstone of Britain, according to Sir Archibald Geikie, "consists of two subdivisions, the lower of which passes down conformably into the Upper Silurian deposits, the upper shading off in the same manner into the base of the Carboniferous system, while they are separated from each other by an unconformability." The Old Red strata appear to have been deposited in a number of elongated lakes or lagoons, approximately parallel to one another, with a general alignment in a N.E.-S.W. direction. To these areas of deposit Sir A. Geikie has assigned convenient distinctive names.
In Scotland the two divisions of the system are sharply separated by a pronounced unconformability which is probably indicative of a prolonged interval of erosion. In the central valley between the base of the Highlands and the southern uplands lay "Lake Caledonia." Here the lower division is made up of some 20,000 ft. of shallow-water deposits, reddish-brown, yellow and grey sandstones and conglomerates, with occasional "cornstones," and thin limestones. The grey flagstones with shales are almost confined to Forfarshire, and are known as the "Arbroath flags." Interbedded volcanic rocks, andesites, dacites, diabases, with agglomerates and tuffs constitute an important feature, and attain a thickness of 6000 ft. in the Pentland and Ochil hills. A line of old volcanic vents may be traced in a direction roughly parallel to the trend of the great central valley. On the northern side of the Highlands was "Lake Orcadie," presumably much larger than the foregoing lake, though its boundaries are not determinable. It lay over Moray Firth and the east of Ross and Sutherland, and extended from Caithness to the Orkney Islands and S. Shetlands. It may even have stretched across to Norway, where similar rocks are found in Sognefjord and Dalsfjord, and may have had communications with some parts of northern Russia. Very characteristic of this area are the Caithness flags, dark grey and bituminous, which, with the red sandstones and conglomerates at their base, probably attain a thickness of 16,000 ft. The somewhat peculiar fauna of this series led Murchison to class the flags as Middle Devonian. In the Shetland Islands contemporaneous volcanic rocks have been observed. Over the west of Argyllshire lay "Lake Lorne"; here the volcanic rocks predominate, they are intercalated with shallow-water deposits. A similar set of rocks occupy the Cheviot district.
The upper division of the Old Red Sandstone is represented in Shropshire and South Wales by a great series of red rocks, shales, sandstones and marls, some 10,000 ft. thick. They contain few fossils, and no break has yet been found in the series. In Scotland this series was deposited in basins which correspond only partially with those of the earlier period. They are well developed in central Scotland over the lowlands bordering the Moray Firth. Interbedded lavas and tuffs are found in the island of Hoy. An interesting feature of this series is the occurrence of great crowds of fossil fishes in some localities, notably at Dura Den in Fife. In the north of England this series rests unconformably upon the Lower Old Red and the Silurian.
Flanking the Silurian high ground of Cumberland and Westmorland, and also in the Lammermuir hills and in Flint and Anglesey, a brecciated conglomerate, presenting many of the characters of a glacial deposit in places, has often been classed with the Old Red Sandstone, but in parts, at least, it is more likely to belong to the base of the Carboniferous system. In Ireland the lower division appears to be represented by the Dingle beds and Glengariff grits, while the Kerry rocks and the Kiltorcan beds of Cork are the equivalents of the upper division. Rocks of Old Red type, both lower and upper, are found in Spitzbergen and in Bear Island. In New Brunswick and Nova Scotia the Old Red facies is extensively developed. The Gaspé sandstones have been estimated at 7036 ft. thick. In parts of western Russia Old Red Sandstone fossils are found in beds intercalated with others containing marine fauna of the Devonian facies.
Devonian and Old Red Sandstone Faunas.
The two types of sediment formed during this period—the marine Devonian and the lagoonal Old Red Sandstone—representing as they do two different but essentially contemporaneous phases of physical condition, are occupied by two strikingly dissimilar faunas. Doubtless at all times there were regions of the earth that were marked off no less clearly from the normal marine conditions of which we have records; but this period is the earliest in which these variations of environment are made obvious. In some respects the faunal break between the older Silurian below and the younger Carboniferous above is not strongly marked; and in certain areas a very close relationship can be shown to exist between the older Devonian and the former, and the younger Devonian and the latter. Nevertheless, taken as a whole, the life of this period bears a distinct stamp of individuality.
The two most prominent features of the Devonian seas are presented by corals and brachiopods. The corals were abundant individually and varied in form; and they are so distinctive of the period that no Devonian species has yet been found either in the Silurian or in the Carboniferous. They built reefs, as in the present day, and contributed to the formation of limestone masses in Devonshire, on the continent of Europe and in North America. Rugose and tabulate forms prevailed; among the former the cyathophyllids (Cyathophyllum) were important, Phillipsastraea, Zaphrentis, Acervularia and the curious Calceola (sandalina), an operculate genus which has given palaeontologists much trouble in its diagnosis, for it has been regarded as a pelecypod (hippurite) and a brachiopod. The tabulate corals were represented by Favosites, Michelinia, Pleurodictyum, Fistulipora, Pachypora and others. Heliolites and Plasmopora represent the alcyonarians. Stromatoporoids were important reef builders. A well-known fossil is Receptaculites, a genus to which it has been difficult to assign a definite place; it has been thought to be a sponge, it may be a calcareous alga, or a curious representative of the foraminifera.
In the Devonian period the brachiopods reached the climax of their development: they compose three-quarters of the known fauna, and more than 1100 species have been described. Changes were taking place from the beginning of the period in the relative importance of genera; several Silurian forms dropped out, and new types were coming in. A noticeable feature was the development of broad-winged shells in the genus Spirifer, other spiriferids were Ambocoelia, Uncites, Verneuilia. Orthids and pentamerids were waning in importance, while the productids (Productella, Chonetes, Strophalosia) were increasing. The strophomenids were still flourishing, represented by the genera Leptaena, Stropheodonta, Kayserella, and others. The ancient Lingula, along with Crania and Orbiculoidea, occur among the inarticulate forms. Another long-lived and wide-ranging species is Atrypa reticularis. The athyrids were very numerous (Athyris, Retzia, Merista, Meristella, Kayserina, &c.); and the rhynchonellids were well represented by Pugnax, Hypothyris, and several other genera. The important group of terebratulids appears in this system; amongst them Stringocephalus is an eminently characteristic Devonian brachiopod; others are Dielasma, Cryptonella, Rensselaeria and Oriskania.
The pelecypod molluscs were represented by Pterinea, abundant in the lower members along with other large-winged forms, and by Cucullella, Buchiola and Curtonotus in the upper members of the system. Other genera are Actinodesma, Cardiola, Nucula, Megalodon, Aviculopecten, &c. Gasteropods were becoming more important, but the simple capulid forms prevailed: Platyceras (Capulus), Straparollus, Pleurotomaria, Murchisonia, Macrocheilina, Euomphalus. Among the pteropods, Tentaculites was very abundant in some quarters; others were Conularia and Styliolina. In the Devonian period the cephalopods began to make a distinct advance in numbers, and in development. The goniatites appear with the genera Anarcestes, Agoniatites, Tornoceras, Bactrites and others; and in the upper strata the clymenoids, forerunners of the later ammonoids, began to take definite shape. While several new nautiloids (Homaloceras, Ryticeras, &c.) made their appearance several of the older genera still lived on (Orthoceras, Poterioceras, Actinoceras).
Crinoids were very abundant in some parts of the Devonian sea, though they were relatively scarce in others; they include the genera Melocrinus, Haplocrinus, Cupressocrinus, Calceocrinus and Eleuthrocrinus. The cystideans were falling off (Proteocystis, Tiaracrinus), but blastoids were in the ascendant (Nucleocrinus, Codaster, &c.). Both brittle-stars, Ophiura, Palaeophiura, Eugaster, and true starfishes, Palaeaster, Aspidosoma, were present, as well as urchins (Lepidocentrus).
When we turn to the crustaceans we have to deal with two distinct assemblages, one purely marine, trilobitic, the other mainly lacustrine or lagoonal with a eurypteridian facies. The trilobites had already begun to decline in importance, and as happens not infrequently with degenerating races of beasts and men, they began to develop strange eccentricities of ornamentation in some of their genera. A number of Silurian genera lived on into the Devonian period, and some gradually developed into new and distinctive forms; such were Proëtus, Harpes, Cheirurus, Bronteus and others. Distinct species of Phacops mark the Lower and Upper Devonian respectively, while the genus Dalmania (Odontochile) was represented by species with an almost world-wide range. The Ostracod Entomis (Cypridina) was extremely abundant in places—Cypridinen-Schiefer—while the true Cypridina was also present along with Beyrichia, Leperditia, &c. The Phyllocarids, Echinocaris, Eleuthrocaris, Tropidocaris, are common in the United States. It is in the Old Red Sandstone that the eurypterids are best preserved; foremost among these was Pterygotus; P. anglicus has been found in Scotland with a length of nearly 6 ft.; Eurypterus, Slimonia, Stylonurus were other genera.
Insects appear well developed, including both orthopterous and neuropterous forms, in the New Brunswick rocks. Mr Scudder believed he had obtained a specimen of Orthoptera in which a stridulating organ was present. A species of Ephemera, allied to the modern may-fly, had a spread of wing extending to 5 in. In the Scottish Old Red Sandstone myriapods, Kampecaris and Archidesmus, have been described; they are somewhat simpler than more recent forms, each segment being separate, and supplied with only one pair of walking legs. Spiders and scorpions also lived upon the land.
The great number of fish remains in the Devonian and Old Red strata, coupled with the truly remarkable characters possessed by some of the forms, has caused the period to be described as the "age of fishes." As in the case of the crustaceans, referred to above, we find one assemblage more or less peculiar to the freshwater or brackish conditions of the Old Red, and another characteristic of the marine Devonian; on the whole the former is the richer in variety, but there seems little doubt that quite a number of genera were capable of living in either environment, whatever may have been the real condition of the Old Red waters. Foremost in interest are the curious ostracoderms, a remarkable group of creatures possessing many of the characteristics of fishes, but more probably belonging to a distinct class of organisms, which appears to link the vertebrates with the arthropods. They had come into existence late in Silurian times; but it is in the Old Red strata that their remains are most fully preserved. They were abundant in the fresh or brackish waters of Scotland, England, Wales, Russia and Canada, and are represented by such forms as Pteraspis, Cephalaspis, Cyathaspis, Tremataspis, Bothriolepis and Pterichthys.
In the lower members of the Old Red series Dipterus, and in the upper members Phaneropleuron, represented the dipnoid lung-fishes; and it is of extreme interest to note that a few of these curious forms still survive in the African Protopterus, the Australian Ceratodus and the South American Lepidosiren,—all freshwater fishes. Distantly related to the lung-fishes were the singular arthrodirans, a group possessing the unusual faculty of moving the head in a vertical plane. These comprise the wide-ranging Coccosteus with Homosteus and Dinichthys, the largest fish of the period. The latter probably reached 20 ft. in length; it was armed with exceedingly powerful jaws provided with turtle-like beaks. Sharks were fairly prominent denizens of the sea; some were armed with cutting teeth, others with crushing dental plates; and although they were on the whole marine fishes, they were evidently able to live in fresher waters, like some of their modern representatives, for their remains, mostly teeth and large dermal spines, are found both in the Devonian and Old Red rocks. Mesacanthus, Diplacanthus, Climatius, Cheiracanthus are characteristic genera. The crossopterygians, ganoids with a scaly lobe in the centre of the fins, were represented by Holoptychius and Glyptopomus in the Upper Old Red, and by such genera as Diplopterus, Osteolepis, Gyroptychius in the lower division. The Polypterus of the Nile and Calamoichthys of South Africa are the modern exemplars of this group. Cheirolepis, found in the Old Red of Scotland and Canada, is the only Devonian representative of the actinopterygian fishes. The cyclostome fishes have, so far, been discovered only in Scotland, in the tiny Palaeospondylus. Amphibian remains have been found in the Devonian of Belgium; and footprints supposed to belong to a creature of the same class (Thinopus antiquus) have been described by Professor Marsh from the Chemung formation of Pennsylvania.
Plant Life.—In the lacustrine deposits of the Old Red Sandstone we find the earliest well-defined assemblage of terrestrial plants. In some regions so abundant are the vegetable remains that in places they form thin seams of veritable coal. These plants evidently flourished around the shores of the lakes and lagoons in which their remains were buried along with the other forms of life. Lycopods and ferns were the predominant types; and it is important to notice that both groups were already highly developed. The ferns include the genera Sphenopteris, Megalopteris, Archaeopteris, Neuropteris. Among the Lycopods are Lycopodites, Psilophyton, Lepidodendron. Modern horsetails are represented by Calamocladus, Asterocalamites, Annularia. Of great interest are the genera Cordaites, Araucarioxylon, &c., which were synthetic types, uniting in some degree the Coniferae and the Cycadofilicales. With the exception of obscure markings, aquatic plants are not so well represented as might have been expected; Parka, a common fossil, has been regarded as a water plant with a creeping stem and two kinds of sporangia in sessile sporocarps.
Physical Conditions, &c.—Perhaps the most striking fact that is brought out by a study of the Devonian rocks and their fossils is the gradual transgression of the sea over the land, which took place quietly in every quarter of the globe shortly after the beginning of the period. While in most places the Lower Devonian sediments succeed the Silurian formations in a perfectly conformable manner, the Middle and Upper divisions, on account of this encroachment of the sea, rest unconformably upon the older rocks, the Lower division being unrepresented. This is true over the greater part of South America, so far as our limited knowledge goes, in much of the western side of North America, in western Russia, in Thuringia and other parts of central Europe. Of the distribution of land and sea and the position of the coast lines in Devonian times we can state nothing with precision. The known deposits all point to shallow waters of epicontinental seas; no abyssal formations have been recognized. E. Kayser has pointed out the probability of a Eurasian sea province extending through Europe towards the east, across north and central Asia towards Manitoba in Canada, and an American sea province embracing the United States, South America and South Africa. At the same time there existed a great North Atlantic land area caused partly by the uplift of the Caledonian range just before the beginning of the period, which stretched across north Europe to eastern Canada; on the fringe of this land the Old Red Sandstone was formed.
In the European area C. Barrois has indicated the existence of three zones of deposition: (1) A northern, Old Red, region, including Great Britain, Scandinavia, European Russia and Spitzbergen; here the land was close at hand; great brackish lagoons prevailed, which communicated more or less directly with the open sea. In European Russia, during its general advance, the sea occasionally gained access to wide areas, only to be driven off again, during pauses in the relative subsidence of the land, when the continued terrigenous sedimentation once more established the lagoonal conditions. These alternating phases were frequently repeated. (2) A middle region, covering Devonshire and Cornwall, the Ardennes, the northern part of the lower Rhenish mountains, and the upper Harz to the Polish Mittelgebirge; here we find evidence of a shallow sea, clastic deposits and a sublittoral fauna. (3) A southern region reaching from Brittany to the south of the Rhenish mountains, lower Harz, Thuringia and Bohemia; here was a deeper sea with a more pelagic fauna. It must be borne in mind that the above-mentioned regions are intended to refer to the time when the extension of the Devonian sea was near its maximum. In the case of North America it has been shown that in early and middle Devonian time more or less distinct faunas invaded the continent from five different centres, viz. the Helderberg, the Oriskany, the Onondaga, the southern Hamilton and the north-western Hamilton; these reached the interior approximately in the order given.
Towards the close of the period, when the various local faunas had mingled one with another and a more generalized life assemblage had been evolved, we find many forms with a very wide range, indicating great uniformity of conditions. Thus we find identical species of brachiopods inhabiting the Devonian seas of England, France, Belgium, Germany, Russia, southern Asia and China; such are, Hypothyris (Rhynchonella) cuboides, Spirifer disjunctus and others. The fauna of the Calceola shales can be traced from western Europe to Armenia and Siberia; the Stringocephalus limestones are represented in Belgium, England, the Urals and Canada; and the (Gephyroceras) intumescens shales are found in western Europe and in Manitoba.
The Devonian period was one of comparative quietude; no violent crustal movements seem to have taken place, and while some changes of level occurred towards its close in Great Britain, Bohemia and Russia, generally the passage from Devonian to Carboniferous conditions was quite gradual. In later periods these rocks have suffered considerable movement and metamorphism, as in the Harz, Devonshire and Cornwall, and in the Belgian coalfields, where they have frequently been thrust over the younger Carboniferous rocks. Volcanic activity was fairly widespread, particularly during the middle portion of the period. In the Old Red rocks of Scotland there is a great thickness (6000 ft.) of igneous rocks, including diabases and andesitic lavas with agglomerates and tuffs. In Devonshire diabases and tuffs are found in the middle division. In west central Europe volcanic rocks are found at many horizons, the most common rocks are diabases and diabase tuffs, schalstein. Felsitic lavas and tuffs occur in the Middle Devonian of Australia. Contemporaneous igneous rocks are generally absent in the American Devonian, but in Nova Scotia and New Brunswick there appear to be some.
There is little evidence as to the climate of this period, but it is interesting to observe that local glacial conditions may have existed in places, as is suggested by the coarse conglomerate with striated boulders in the upper Old Red of Scotland. On the other hand, the prevalence of reef-building corals points to moderately warm temperatures in the Middle Devonian seas.
The economic products of Devonian rocks are of some importance: in many of the metamorphosed regions veins of tin, lead, copper, iron are exploited, as in Cornwall, Devon, the Harz; in New Zealand, gold veins occur. Anthracite of Devonian age is found in China and a little coal in Germany, while the Upper Devonian is the chief source of oil and gas of western Pennsylvania and south-western New York. In Ontario the middle division is oil-bearing. Black phosphates are worked in central Tennessee, and in England the marls of the "Old Red" are employed for brick-making.