- Key People:
- William Lonsdale
- Robert Kidston
A highly varied invertebrate fauna that originated in the preceding Silurian Period continued in the Devonian, and most ecological niches of shallow and deep marine water were exploited. The remarkable proliferation of primitive fishes, which has given the period the name the “Age of Fishes,” occurred in both fresh and marine waters. Derivation of carnivorous fishes from mud-eating forms occurred early in the period, and tetrapods (four-legged land animals) were derived from fishes near the middle of the period. Also remarkable is the rise to dominance of vascular plants. Though groves of trees must have arisen earlier to provide the widespread plant debris noted in Devonian deposits, the first known evidence of in-place forests dates from the Middle Devonian.
Invertebrates
The Devonian invertebrates are essentially of the type established during the Ordovician Period. In nearshore sandy and silty environments, bivalves, burrowing organisms, brachiopods (lamp shells), and simple corals abounded. In offshore environments free from land detritus, biostromes and bioherms flourished, rich in corals, stromatoporoids (large colonial marine organisms similar to hydrozoans), crinoids, brachiopods, trilobites, gastropods, and other forms. In deeper waters, goniatite ammonites (a form of cephalopod), which were one of the few new groups to appear, were abundant. Surface waters were occupied by small dacryoconarids (a shelled marine invertebrate) and by ostracods (mussel shrimp) later in the period. Among the Protozoa, both Foraminifera and Radiolaria were well represented, and sponges were locally abundant.
The corals and stromatoporoids were extremely important for building reef facies. The limestone-reef and forereef facies and biostromal limestones are known in many areas of the world. The corals include tabulate corals, such as Favosites and Alveolites, but especially rugose corals (horn corals), which have been used to establish correlations. Stromatoporoids (a type of sponge with a layered skeleton composed of calcium carbonates) such as Amphipora were common rock builders in the mid-Devonian of the Northern Hemisphere. The twiglike form of Amphipora produces a “spaghetti” or “vermicelli” rock. Elsewhere, only simple corals are frequently found.
Bryozoans (marine moss animals superficially similar to corals) were especially common in shallow shelf seas of the period. Both stony and netted forms occurred, but only the latter, the fenestellids, became important during the period.
The brachiopods (lamp shells) are a group of marine filter-feeding species that bear a resemblance to clams but are not mollusks. Brachiopods were present in a multitude of diverse forms during the Devonian Period. The spire-bearing spiriferoids were perhaps the most common and have been used as index fossils. Two groups of importance emerged: the loop-bearing terebratulids and the spiny mud-dwelling productids. At the same time, a number of groups became extinct, including various orthids and the pentamerids.
Molluscan groups were well represented. The marine clams (bivalves) diversified greatly during the period, especially in the nearshore environments. The earliest freshwater bivalves appeared in the Late Devonian. The gastropods were well diversified, particularly in calcareous (calcium carbonate or limestone) environments, and became even more diversified in later periods. The Scaphopoda (tusk shells) first appeared in the Devonian Period. Another significant Devonian event was the emergence of the ammonites from their still-extant nautiloid ancestors. In the chambered shell of the ammonites, internal septa create elaborate patterns where they join the outer shell. The complexity of these suture patterns culminated in the ammonites of the Mesozoic Era. From their origin (probably in the Emsian Age) the evolution of goniatite ammonites, as well as other ammonites, allows detailed zonal subdivisions to be established until the end of the Cretaceous Period. Devonian goniatites have been found on all continents except Antarctica.
Among the arthropods, the giant eurypterids (sea scorpions) are found in the Old Red Sandstone facies. Some were predatory carnivores and probably lived on fish. The first insect, most likely a collembolan (apterygote), from a group of wingless insects that feed on leaf litter and soil, has been recorded from the Devonian Period of Russia and other areas of Asia. Ostracods (a type of crustacean) were locally very abundant; benthic (bottom-dwelling) forms occur in continental shelf sea deposits, and planktonic (floating) forms occur in the Upper Devonian, where their remains form widespread ostracod-slate facies. Trilobites were well developed in size (some up to 61 cm, or 24 inches, long), variety, and distribution. Nearly all have clearly established Silurian ancestors. The most common were the phacopids, which exhibit a curious trend toward blindness in the Late Devonian. Almost all the diverse Lower Paleozoic trilobite stocks that entered the period were extinct before the close, and only the proetaceans survived into the Carboniferous Period.
Among echinoderms, the holothureans, asteroids, and ophiuroids are known, but they are rare. Crinoids were abundant, including free-living types with grapnel-shaped anchors. The blastoids diversified considerably, but the cystoids did not survive the period.
Vertebrates
Conodonts (recently recognized as toothlike elements of very primitive eel-like vertebrates) are abundant in many Devonian marine facies. Conodonts had perhaps their greatest diversification during the Late Devonian and are of major importance for the correlation of rock layers. More than 40 conodont zones are recognized within the Devonian, and these provide a high-resolution biostraphic framework for the period.
Many groups of Devonian fishes were heavily armoured, and this has led to their good representation in the fossil record. Fish remains are widespread in the Old Red Sandstone rocks of Europe, especially in the Welsh borderland and Scottish areas of Britain; these are mostly associated with freshwater or estuarine deposits. In other areas marine fishes are known, and some of these, such as Dunkleosteus (Dinichthys) from the Upper Devonian Period of Ohio, U.S., may have reached 9 metres (30 feet) in length.
The earliest fishes, comprising the agnathans, were without jaws and presumably were mud eaters and scavengers. These types are usually called ostracoderms. Some, such as the osteostracan cephalaspids, had broad, platelike armour of varied form; the brain and nerve structures in some of these are well known. The anaspids also were covered with armour in the form of scales. The heterostracans, which include the oldest known fishes, have an anterior armour basically of upper (dorsal) and lower (ventral) plates; Pteraspis is an example. The Early Devonian saw the entry of jawed forms or gnathostomes, and the armoured forms of these, the placoderms, characterize the epoch. The arthrodires, which had a hinged frontal armour in two portions, and the grotesque antiarchs belong here. The close of the Devonian saw the diminution and extinction of most of these groups, but several other groups continued and have a significant later history.
Sharklike fishes, the chondrichthians, have been found in the Middle Devonian. The bony fishes, or osteichthians of current classification, include the climatioid acanthodeans, which had appeared before the period began, but the lungfishes (Dipnoi), the coelacanths, and the rhipidistians made their first appearance during this time. The last group is thought to have given rise to the four-footed amphibians as well as to all other higher groups of vertebrates.