The Devonian is a geologic period and system of the Paleozoic Era spanning from the end
of the Silurian Period, about 419.2 ± 3.2 Mya, to the beginning of the Carboniferous
Period, about 358.9 ± 0.4. It is named after Devon, England, where rocks from this period
were first studied.
The Devonian period experienced the first significant adaptive radiation of terrestrial
life. Free-sporing vascular plants began to spread across dry land, forming extensive
forests which covered the continents. By the middle of the Devonian, several groups of
plants had evolved leaves and true roots, and by the end of the period the first seed-bearing
plants appeared. Various terrestrial arthropods also became well-established. Fish reached
substantial diversity during this time, leading the Devonian to often be dubbed the "Age of
Fish". The first ray-finned and lobe-finned bony fish appeared, while the placoderms began
dominating almost every known aquatic environment. The ancestors of all tetrapods began adapting
to walking on land, their strong pectoral and pelvic fins gradually evolved into legs.
In the oceans, primitive sharks became more numerous than in the Silurian and the late
Ordovician. The first ammonite mollusks appeared. Trilobites, the mollusk-like brachiopods and
the great coral reefs, were still common. The Late Devonian extinction which started
about 375 million years ago, severely affected marine life, killing off all placoderms, and
all trilobites, save for a few species of the order Proetida.
The paleogeography was dominated by the supercontinent of Gondwana to the south, the continent of
Siberia to the north, and the early formation of the small continent of Euramerica in between.
History
The period is named after Devon, a county in southwestern England, where a controversial
argument in the 1830s over the age and structure of the rocks found distributed throughout
the county was eventually resolved by the defining of the Devonian period in the geological
timescale. The Great Devonian Controversy is a classic case of how the foundations of
our present-day geological knowledge and classification of the rock record and geological timescale
was socially as well as scientifically constructed. After a long period of vigorous argument and
counter-argument between the main protagonists of Roderick Murchison with Adam Sedgwick against
Henry de la Beche supported by George Bellas Greenough, Murchison and Sedgwick won the
debate and named the period they proposed as 'The Devonian System'.
While the rock beds that define the start and end of the Devonian period are well identified,
the exact dates are uncertain. According to the International Commission on Stratigraphy,
the Devonian extends from the end of the Silurian Period 419.2 ± 3.2 Mya, to the beginning
of the Carboniferous Period 358.9 ± 0.4 Mya (in North America, the beginning of the Mississippian
subperiod of the Carboniferous. In nineteenth-century texts the Devonian has
been called the "Old Red Age", after the red and brown terrestrial deposits known in the
United Kingdom as the Old Red Sandstone in which early fossil discoveries were found.
Another common term is "Age of the Fishes", referring to the evolution of several major
groups of fish that took place during the period. Older literature on the Anglo-Welsh
basin divides it into the Downtonian, Dittonian, Breconian and Farlovian stages, the latter
three of which are placed in the Devonian. The Devonian has also erroneously been characterized
as a "greenhouse age", due to sampling bias: most of the early Devonian-age discoveries
came from the strata of western Europe and eastern North America, which at the time straddled
the Equator as part of the supercontinent of Euramerica where fossil signatures of widespread
reefs indicate tropical climates that were warm and moderately humid but in fact the
climate in the Devonian differed greatly between epochs and geographic regions. For example,
during the Early Devonian, arid conditions were prevalent through much of the world including
Siberia, Australia, North America, and China, but Africa and South America had a warm temperate
climate. In the Late Devonian, by contrast, arid conditions were less prevalent across
the world and temperate climates were more common.
Subdivisions
The Devonian Period is formally broken into Early, Middle and Late subdivisions. The rocks
corresponding to these epochs are referred to as belonging to the Lower, Middle and Upper
parts of the Devonian System. The Early Devonian lasts from 419.2 ± 2.8 to
393.3 ± 2.5 and begins with the Lochkovian stage, which lasts until the Pragian. This
spans from 410.8 ± 2.8 to 407.6 ± 2.5, and is followed by the Emsian, which lasts
until the Middle Devonian begins, 393.3± 2.7 million years ago. The Middle Devonian comprises
two subdivisions, the Eifelian giving way to the Givetian 387.7± 2.7 million years
ago. During this time the armoured jawless ostracoderm fish were declining in diversity;
the jawed fish were thriving and increasing in diversity in both the oceans and freshwater.
The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive
plants, provided the environment necessary for certain early fish to develop essential
characteristics such as well developed lungs, and the ability to crawl out of the water
and onto the land for short periods of time. Finally, the Late Devonian starts with the
Frasnian, 382.7 ± 2.8 to 372.2 ± 2.5, during which the first forests were taking
shape on land. The first tetrapods appear in the fossil record in the ensuing Famennian
subdivision, the beginning and end of which are marked with extinction events. This lasted
until the end of the Devonian, 358.9± 2.5 million years ago.
Climate The Devonian was a relatively warm period,
and probably lacked any glaciers. The temperature gradient from the equator to the poles was
not as large as it today. The weather was also very arid, mostly along the equator where
it was the driest. Reconstruction of tropical sea surface temperature from conodont apatite
implies an average value of 30 °C in the Early Devonian. CO
2 levels dropped steeply throughout the Devonian period as the burial of the newly evolved
forests drew carbon out of the atmosphere into sediments; this may be reflected by a
Mid-Devonian cooling of around 5 °C. The Late Devonian warmed to levels equivalent
to the Early Devonian; while there is no corresponding increase in CO
2 concentrations, continental weathering increases; further, a range of evidence, such as plant
distribution, points to Late Devonian warming. The climate would have affected the dominant
organisms in reefs; microbes would have been the main reef-forming organisms in warm periods,
with corals and stromatoporoid sponges taking the dominant role in cooler times. The warming
at the end of the Devonian may even have contributed to the extinction of the stromatoporoids.
Paleogeography
The Devonian period was a time of great tectonic activity, as Euramerica and Gondwana drew
closer together. The continent Euramerica was created in the
early Devonian by the collision of Laurentia and Baltica, which rotated into the natural
dry zone along the Tropic of Capricorn, which is formed as much in Paleozoic times as nowadays
by the convergence of two great air-masses, the Hadley cell and the Ferrel cell. In these
near-deserts, the Old Red Sandstone sedimentary beds formed, made red by the oxidized iron
characteristic of drought conditions. Near the equator, the plate of Euramerica
and Gondwana were starting to meet, beginning the early stages of assembling Pangaea. This
activity further raised the northern Appalachian Mountains and formed the Caledonian Mountains
in Great Britain and Scandinavia. The west coast of Devonian North America,
by contrast, was a passive margin with deep silty embayments, river deltas and estuaries,
in today's Idaho and Nevada; an approaching volcanic island arc reached the steep slope
of the continental shelf in Late Devonian times and began to uplift deep water deposits,
a collision that was the prelude to the mountain-building episode of Mississippian times called the
Antler orogeny. Sea levels were high worldwide, and much of
the land lay under shallow seas, where tropical reef organisms lived. The deep, enormous Panthalassa
covered the rest of the planet. Other minor oceans were Paleo-Tethys, Proto-Tethys, Rheic
Ocean, and Ural Ocean. Biota
Marine biota Sea levels in the Devonian were generally
high. Marine faunas continued to be dominated by bryozoa, diverse and abundant brachiopods,
the enigmatic hederelloids, microconchids and corals. Lily-like crinoids were abundant,
and trilobites were still fairly common. Among vertebrates, jaw-less armored fish declined
in diversity, while the jawed fish simultaneously increased in both the sea and fresh water.
Armored placoderms were numerous during the lower stages of the Devonian Period and became
extinct in the Late Devonian, perhaps because of competition for food against the other
fish species. Early cartilaginous and bony fishes also become diverse and played a large
role within the Devonian seas. The first abundant genus of shark, Cladoselache, appeared in
the oceans during the Devonian Period. The great diversity of fish around at the time,
have led to the Devonian being given the name "The Age of Fish" in popular culture.
The first ammonites also appeared during or slightly before the early Devonian Period
around 400 Mya.
Reefs A now dry barrier reef, located in present
day Kimberley Basin of northwest Australia, once extended a thousand kilometers, fringing
a Devonian continent. Reefs in general are built by various carbonate-secreting organisms
that have the ability to erect wave-resistant frameworks close to sea level. The main contributors
of the Devonian reefs were unlike modern reefs, which are constructed mainly by corals and
calcareous algae. They were composed of calcareous algae and coral-like stromatoporoids, and
tabulate and rugose corals, in that order of importance.
Terrestrial biota By the Devonian Period, life was well underway
in its colonization of the land. The moss forests and bacterial and algal mats of the
Silurian were joined early in the period by primitive rooted plants that created the first
stable soils and harbored arthropods like mites, scorpions and myriapods. Also the first
possible fossils of insects appeared around 416 Mya in the Early Devonian. The first tetrapods,
evolving from lobe-finned fish, appeared in the coastal water no later than middle Devonian,
and gave rise to the first Amphibians. The greening of land
Early Devonian plants did not have roots or leaves like the plants most common today and
many had no vascular tissue at all. They probably spread largely by vegetative growth, and did
not grow much more than a few centimeters tall. By far the largest land organism was
Prototaxites, the fruiting body of an enormous fungus that stood more than 8 meters tall,
towering over the low, carpet-like vegetation. By the Middle Devonian, shrub-like forests
of primitive plants existed: lycophytes, horsetails, ferns, and progymnosperms had evolved. Most
of these plants had true roots and leaves, and many were quite tall. The earliest known
trees, from the genus Wattieza, appeared in the Late Devonian around 385 Ma. In the Late
Devonian, the tree-like ancestral Progymnosperm Archaeopteris which had conifer-like true
wood and fern-like foliage and the cladoxylopsids grew. These are the oldest known trees of
the world's first forests. By the end of the Devonian, the first seed-forming plants had
appeared. This rapid appearance of so many plant groups and growth forms has been called
the "Devonian Explosion". The 'greening' of the continents acted as
a carbon dioxide sink, and atmospheric levels of this greenhouse gas may have dropped. This
may have cooled the climate and led to a massive extinction event. See Late Devonian extinction.
Animals and the first soils Primitive arthropods co-evolved with this
diversified terrestrial vegetation structure. The evolving co-dependence of insects and
seed-plants that characterizes a recognizably modern world had its genesis in the Late Devonian
period. The development of soils and plant root systems probably led to changes in the
speed and pattern of erosion and sediment deposition. The rapid evolution of a terrestrial
ecosystem containing copious animals opened the way for the first vertebrates to seek
out a terrestrial living. By the end of the Devonian, arthropods were solidly established
on the land. Late Devonian extinction
A major extinction occurred at the beginning of the last phase of the Devonian period,
the Famennian faunal stage,, about 372.2 ± 1.6 Mya, when all the fossil agnathan fishes,
save for the psammosteid heterostracans, suddenly disappeared. A second strong pulse closed
the Devonian period. The Late Devonian extinction was one of five major extinction events in
the history of the Earth's biota, more drastic than the familiar extinction event that closed
the Cretaceous. The Devonian extinction crisis primarily affected
the marine community, and selectively affected shallow warm-water organisms rather than cool-water
organisms. The most important group to be affected by this extinction event were the
reef-builders of the great Devonian reef-systems. Amongst the severely affected marine groups
were the brachiopods, trilobites, ammonites, conodonts, and acritarchs, as well as jawless
fish, and all placoderms. Land plants as well as freshwater species, such as our tetrapod
ancestors, were relatively unaffected by the Late Devonian extinction event.
The reasons for the Late Devonian extinctions are still unknown, and all explanations remain
speculative. Canadian paleontologist Digby McLaren suggested in 1969 that the Devonian
extinction events were caused by an asteroid impact. However, while there were Late Devonian
collision events, little evidence supports the existence of a large enough Devonian crater.
See also Geologic timescale
Phacops rana: a Devonian trilobite. List of fossil sites
Falls of the Ohio State Park USA, Indiana. One of the largest exposed Devonian fossil
beds in the world. References
References Ogg, Jim; June, 2004, Overview of Global Boundary
Stratotype Sections and Points http:www.stratigraphy.org/gssp.htm Accessed April 30, 2006.
Palaeos Age of Fishes Museum
External links The Devonian times - an excellent and frequently
updated resource focussing on the Devonian period
UC Berkeley site introduces the Devonian. "International Commission on Stratigraphy".
Geologic Time Scale 2004. Retrieved September 19, 2005.
Examples of Devonian Fossils