Devonian

The Devonian ( ) is a geologic period and system of the Paleozoic, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after

Devon

, England, where rocks from this period were first studied.

The first significant adaptive radiation of life on dry land occurred during the Devonian. Free-sporing vascular plants began to spread across dry land, forming extensive

forest

s 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. The arthropod groups of myriapods, arachnids and hexapods also became well-established early on in this period, after having started their expansion to land at least from the

Ordovician

period.

Fish

reached substantial diversity during this time, leading the Devonian to often be dubbed the Age of Fishes. The placoderms began dominating almost every known aquatic environment. The ancestors of all four-limbed vertebrates (tetrapods) began adapting to walk on land, as their strong pectoral and pelvic fins gradually evolved into legs, though they were not fully established until the Late Carboniferous. In the oceans, primitive

shark

s became more numerous than in the Silurian and Late Ordovician.

The first

ammonites

, a subclass of molluscs, appeared.

Trilobite

s, the mollusc-like brachiopods, and the great

coral reef

s were still common. The Late Devonian extinction which started about 375 million years ago severely affected marine life, killing off all placodermi, and all trilobites, save for a few species of the order Proetida.

The palaeogeography was dominated by the supercontinent of

Gondwana

to the south, the small continent of Siberia to the north, and the early formation of the medium-sized 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 definition of the Devonian Period in the geological timescale. The Great Devonian Controversy was 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 Mya, to the beginning of the Carboniferous Mya – in

North America

, at 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 characterised 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

climate

s that were warm and moderately humid. In fact the climate in the Devonian differed greatly during its

epochs

and between 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 those epochs are referred to as belonging to the Lower, Middle and Upper parts of the Devonian System.

;Early Devonian
The Early Devonian lasted from and began with the Lochkovian Stage , which was followed by the Pragian from and then by the Emsian, which lasted until the Middle Devonian began, .
During this time, the first

ammonoids

appeared, descending from bactritoid nautiloids. Ammonoids during this time period were simple and differed little from their nautiloid counterparts. These ammonoids belong to the order Agoniatitida, which in later epochs evolved to new ammonoid orders, for example Goniatitida and Clymeniida. This class of cephalopod molluscs would dominate the marine fauna until the beginning of the Mesozoic Era.

;Middle Devonian
The Middle Devonian comprised two subdivisions: first the Eifelian, which then gave way to the Givetian . During this time the jawless agnathan fishes began to decline in diversity in freshwater and marine environments partly due to drastic environmental changes and partly due to the increasing competition, predation, and diversity of

jawed fishes

. The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop such essential characteristics as well developed lungs, and the ability to crawl out of the water and onto the land for short periods of time.

;Late Devonian
Finally, the Late Devonian started with the Frasnian, , during which the first forests took shape on land. The first tetrapods appeared 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, .

Climate

The Devonian was a relatively warm period, and probably lacked any

glacier

s. The temperature gradient from the equator to the poles was not as large as it is 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 in the Early Devonian. levels dropped steeply throughout the Devonian Period. The newly evolved forests drew carbon out of the atmosphere, which were then buried into sediments. This may be reflected by a Mid-Devonian cooling of around . The Late Devonian warmed to levels equivalent to the Early Devonian; while there is no corresponding increase in concentrations, continental weathering increases (as predicted by warmer temperatures); further, a range of evidence, such as plant distribution, points to a Late Devonian warming. The climate would have affected the dominant organisms in

reef

s; 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 (or Laurussia) 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 oxidised iron (

hematite

) characteristic of drought conditions.

Near the

equator

, the plate of Euramerica and Gondwana were starting to meet, beginning the early stages of the assembling of

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, found today in

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 at the beginning of the Carboniferous 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 (the "universal ocean") covered the rest of the

planet

. Other minor oceans were the Paleo-Tethys Ocean, Proto-Tethys Ocean, Rheic Ocean, and Ural Ocean (which was closed during the collision with Siberia and Baltica).

During the Devonian, Chaitenia, an

island arc

, accreted to Patagonia.

Life

Marine biota

Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by

bryozoa

, diverse and abundant brachiopods, the enigmatic hederellids, microconchids and

coral

s. Lily-like

crinoid

s (animals, their resemblance to flowers notwithstanding) were abundant, and

trilobite

s were still fairly common. Among vertebrates, jawless armored fish (

ostracoderm

s) declined in diversity, while the jawed fish (gnathostomes) 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 (

Chondrichthyes

) and bony fishes (

Osteichthyes

) 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 has 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 , fringing a Devonian continent. Reefs in general are built by various

carbonate

-secreting organisms that have the ability to erect wave-resistant structures close to sea level. Although modern reefs are constructed mainly by corals and calcareous

algae

, Devonian reefs were either microbial reefs built up mostly by autotrophic

cyanobacteria

, or coral-stromatoporoid reefs built up by coral-like stromatoporoids and tabulate and rugose corals. Microbial reefs dominated under the warmer conditions of the early and late Devonian, while coral-stromatoporoid reefs dominated during the cooler middle Devonian.

Terrestrial biota

By the Devonian Period, life was well underway in its colonisation of the land. The

moss

forests and

bacteria

l and algal mats of the Silurian were joined early in the period by primitive rooted plants that created the first stable

soil

s and harbored arthropods like mites,

scorpion

s, trigonotarbids and

myriapods

(although arthropods appeared on land much earlier than in the Early Devonian and the existence of fossils such as ''

Protichnites

'' suggest that amphibious arthropods may have appeared as early as the Cambrian). By far the largest land organism at the beginning of this period was the enigmatic ''Prototaxites'', which was possibly the fruiting body of an enormous fungus, rolled liverwort mat, or another organism of uncertain affinities that stood more than tall, and towered over the low, carpet-like vegetation during the early part of the Devonian. Also the first possible fossils of insects appeared around 416 Mya, in the Early Devonian. Evidence for the earliest tetrapods takes the form of trace fossils in shallow lagoon environments within a marine carbonate platform / shelf during the Middle Devonian, although these traces have been questioned and an interpretation as fish feeding traces (''Piscichnus'') has been advanced.

The greening of land

Many List of Early Devonian land plants, Early Devonian plants did not have true roots or leaves like extant plants although vascular tissue is observed in many of those plants. Some of the early land plants such as ''Drepanophycus'' likely spread by vegetative growth and spores. The earliest land plants such as ''Cooksonia'' consisted of leafless, Dichotomous branching, dichotomous axes and terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall. Fossils of ''Armoricaphyton, Armoricaphyton chateaupannense'', about 400 million years old, represent the oldest known plants with woody tissue. By the Middle Devonian, shrub-like forests of primitive plants existed: Lycopodiophyta, lycophytes, Equisetales, horsetails, ferns, and progymnosperms had evolved. Most of these plants had true roots and leaves, and many were quite tall. The earliest-known trees appeared in the Middle Devonian. These included a lineage of lycopods and another arborescent, woody vascular plant, the Cladoxylopsida, cladoxylopsids. These tracheophytes were able to grow to large size on dry land because they had evolved the ability to biosynthesize lignin, which gave them physical rigidity and improved the effectiveness of their vascular system while giving them resistance to pathogens and herbivores. 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 sink, and Atmosphere of Earth, atmospheric concentrations of carbon dioxide 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 characterised a recognisably modern world had its genesis in the Late Devonian Epoch. 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 that contained 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.

Gallery

Image:D_Terrelli.png, ''Dunkleosteus'', one of the largest armoured fish ever to roam the planet, lived during the Late Devonian
Image:Eastmanosteus pustulosus.jpg, Lower jaw of ''Eastmanosteus pustulosus'' from the Middle Devonian of Wisconsin
Image:Onychodus.jpg, Tooth of the lobe-finned fish ''Onychodus'' from the Middle Devonian of Wisconsin
Image:Devonianfishes ntm 1905 smit 1929.gif, Early shark ''

Cladoselache

'', several Sarcopterygii, lobe-finned fishes, including ''Eusthenopteron'' that was an early marine tetrapod, and the placoderm ''Bothriolepis'' in a painting from 1905
Image:Melocrinites nodosus spinosus.jpg, ''Melocrinites nodosus spinosus'', a spiny, stalked

crinoid

from the Middle Devonian of Wisconsin
Image:PhacopidDevonian.jpg, Enrolled phacopid

trilobite

from the Devonian of Ohio
Image:AuloporaDevonianSilicaShale.jpg, The common tabulate coral ''Aulopora'' from the Middle Devonian of Ohio – view of colony encrusting a brachiopod valve
Image:Tropidoleptus carinatus.jpg, ''Tropidoleptus carinatus'', an orthid brachiopod from the Middle Devonian of New York
Image:Pleurodictyum americanum Kashong.jpg, ''Pleurodictyum, Pleurodictyum americanum'', Kashong Shale, Middle Devonian of New York
Image:HederelloidSEM.jpg, SEM image of a hederelloid from the Devonian of Michigan (largest tube diameter is 0.75 mm)
Image:HederellaOH3.jpg, Devonian spiriferid brachiopod from Ohio which served as a host substrate for a colony of hederelloids

Late Devonian extinction

The Late Devonian extinction is not a single event, but rather is a series of pulsed extinctions at the Givetian-Frasnian boundary, the Frasnian-Famennian boundary, and the Devonian-Carboniferous boundary. Together, these are considered one of the "Big Five" mass extinctions in Earth's history. 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, and acritarchs, and the world saw the disappearance of an estimated 96% of vertebrates like conodonts and Osteichthyes, bony fishes, and all of the ostracoderms and placoderms. Land plants as well as freshwater species, such as our tetrapod ancestors, were relatively unaffected by the Late Devonian extinction event (there is a counterargument that the Devonian extinctions nearly wiped out the tetrapods).

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 (see the Alamo bolide impact), little evidence supports the existence of a large enough Devonian crater.

See also

* Celsius Bjerg Formation - in Greenland, with fossils dating back to the Devonian Period.
* , Indiana, United States. One of the largest exposed Devonian fossil beds in the world.
*
*
* ''(with link directory)''
* , a Devonian trilobite
;Categories:
*

Notes

References

External links

*
* – site introduces the Devonian
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{{Authority control

Devonian,
Geological periods
Geology of Devon, .