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The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
, Mya. It is named after Devon, England, where rocks from this period were first studied. The first significant
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic in ...
of life on dry land occurred during the Devonian. 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. The
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, o ...
groups of
myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
,
arachnids Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaro ...
and hexapods also became well-established early in this period, after starting their expansion to land at least from the
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...
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 (
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsid ...
s) 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 Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effe ...
. In the oceans, primitive sharks became more numerous than in the Silurian and Late Ordovician. The first
ammonites Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefis ...
, a subclass of molluscs, appeared. Trilobites, the mollusc-like
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, wh ...
s, and the great coral reefs were still common. The
Late Devonian extinction The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major ex ...
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 Proetida is an order of trilobite that lived from the Ordovician to the Permian. It was the last order of trilobite to go extinct, finally dying out in the Permian-Triassic extinction event. Description These typically small trilobites resem ...
. Devonian palaeogeography was dominated by the supercontinent of Gondwana to the south, the small continent of Siberia to the north, and the medium-sized continent of
Laurussia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
to the east. Major tectonic events include the closure of the
Rheic Ocean The Rheic Ocean was an ocean which separated two major palaeocontinents, Gondwana and Laurussia (Laurentia-Baltica-Avalonia). One of the principal oceans of the Palaeozoic, its sutures today stretch from Mexico to Turkey and its closure resulted ...
, the separation of
South China South China () is a geographical and cultural region that covers the southernmost part of China. Its precise meaning varies with context. A notable feature of South China in comparison to the rest of China is that most of its citizens are not n ...
from Gondwana, and the resulting expansion of the
Paleo-Tethys Ocean The Paleo-Tethys or Palaeo-Tethys Ocean was an ocean located along the northern margin of the paleocontinent Gondwana that started to open during the Middle Cambrian, grew throughout the Paleozoic, and finally closed during the Late Triassic; exi ...
. The Devonian experienced several major mountain-building events as Laurussia and Gondwana approached; these include the Acadian Orogeny in North America and the beginning of the Variscan Orogeny in Europe. These early collisions preceded the formation of
Pangaea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
in the Late Paleozoic.


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 The Great Devonian Controversy began in 1834 when Roderick Murchison disagreed with Henry De la Beche as to the dating of certain petrified plants found in coals in the Greywacke stratum in North Devon, England. De La Beche was claiming that ...
was a long period of vigorous argument and counter-argument between the main protagonists of Roderick Murchison with
Adam Sedgwick Adam Sedgwick (; 22 March 1785 – 27 January 1873) was a British geologist and Anglican priest, one of the founders of modern geology. He proposed the Cambrian and Devonian period of the geological timescale. Based on work which he did on W ...
against
Henry De la Beche Sir Henry Thomas De la Beche KCB, FRS (10 February 179613 April 1855) was an English geologist and palaeontologist, the first director of the Geological Survey of Great Britain, who helped pioneer early geological survey methods. He was the f ...
supported by
George Bellas Greenough George Bellas Greenough FRS FGS (18 January 1778 – 2 April 1855) was a pioneering English geologist. He is best known as a synthesizer of geology rather than as an original researcher. Trained as a lawyer, he was a talented speaker and his ...
. 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 The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
Mya – in North America, at the beginning of the Mississippian subperiod of the Carboniferous. In 19th 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 The Old Red Sandstone is an assemblage of rocks in the North Atlantic region largely of Devonian age. It extends in the east across Great Britain, Ireland and Norway, and in the west along the northeastern seaboard of North America. It also exte ...
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 In statistics, sampling bias is a bias in which a sample is collected in such a way that some members of the intended population have a lower or higher sampling probability than others. It results in a biased sample of a population (or non-human f ...
: 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 Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
s that were warm and moderately humid. In fact the climate in the Devonian differed greatly during its
epochs In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
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 The Lochkovian is one of three faunal stages in the Early Devonian Epoch. It lasted from 419.2 ± 3.2 million years ago to 410.8 ± 2.8 million years ago. It marked the beginning of the Devonian Period, and was followed by the Pragian Stage. It is ...
Stage , which was followed by the
Pragian The Pragian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.8 ± 2.8 million years ago to 407.6 ± 2.8 million years ago. It was preceded by the Lochkovian Stage and followed by the Emsian Stage. The most important La ...
from and then by the
Emsian The Emsian is one of three faunal stages in the Early Devonian Epoch. It lasted from 407.6 ± 2.6 million years ago to 393.3 ± 1.2 million years ago. It was preceded by the Pragian Stage and followed by the Eifelian Stage. It is named after the ...
, which lasted until the Middle Devonian began, . During this time, the first
ammonoids Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish ...
appeared, descending from bactritoid
nautiloids Nautiloids are a group of marine cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and speciose, with over 2,500 recorded species. ...
. Ammonoids during this time period were simple and differed little from their nautiloid counterparts. These ammonoids belong to the order
Agoniatitida Agoniatitida, also known as the Anarcestida, is the ancestral order within the cephalopod subclass Ammonoidea originating from bactritoid nautiloids, that lived in what would become Africa, Asia, Australia, Europe, and North America during th ...
, which in later epochs evolved to new ammonoid orders, for example Goniatitida and Clymeniida. This class of
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, a ...
molluscs would dominate the marine fauna until the beginning of the Mesozoic Era. ;Middle Devonian The Middle Devonian comprised two subdivisions: first the
Eifelian The Eifelian is the first of two faunal stages in the Middle Devonian Epoch. It lasted from 393.3 ± 1.2 million years ago to 387.7 ± 0.8 million years ago. It was preceded by the Emsian Stage and followed by the Givetian Stage. North American ...
, which then gave way to the Givetian . During this time the jawless
agnathan Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostome ...
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 Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...
. 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 The Frasnian is one of two faunal stages in the Late Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Givetian Stage and followed by the Famennian Stage. Major reef-building was under way during th ...
, , during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing
Famennian The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used ...
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 glaciers for much of the period. 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 Conodonts (Greek ''kōnos'', " cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, whic ...
apatite Apatite is a group of phosphate minerals, usually hydroxyapatite, fluorapatite and chlorapatite, with high concentrations of OH−, F− and Cl− ions, respectively, in the crystal. The formula of the admixture of the three most common ...
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 reefs; microbes would have been the main reef-forming organisms in warm periods, with corals and
stromatoporoid Stromatoporoidea is an extinct clade of sea sponges common in the fossil record from the Ordovician through the Devonian. They were especially abundant and important reef-formers in the Silurian and most of the Devonian.Stock, C.W. 2001, Stromat ...
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. At the terminus of the Devonian, Earth rapidly cooled into an icehouse, marking the beginning of the Late Palaeozoic Ice Age.


Paleogeography

The Devonian world involved many continents and ocean basins of various sizes. The largest continent, Gondwana, was located entirely within the Southern Hemisphere. It corresponds to modern day South America,
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
, Australia,
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
, and India, as well as minor components of North America and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
. The second-largest continent, Laurussia, was northwest of Gondwana, and corresponds to much of modern-day North America and Europe. Various smaller continents, microcontinents, and terranes were present east of Laurussia and north of Gondwana, corresponding to parts of Europe and Asia. The Devonian Period was a time of great tectonic activity, as the major continents of
Laurussia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
and Gondwana drew closer together. Sea levels were high worldwide, and much of the land lay under shallow seas, where tropical reef organisms lived. The enormous "world ocean", Panthalassa, occupied much of the Northern Hemisphere as well as wide swathes east of Gondwana and west of Laurussia. Other minor oceans were the
Paleo-Tethys Ocean The Paleo-Tethys or Palaeo-Tethys Ocean was an ocean located along the northern margin of the paleocontinent Gondwana that started to open during the Middle Cambrian, grew throughout the Paleozoic, and finally closed during the Late Triassic; exi ...
and
Rheic Ocean The Rheic Ocean was an ocean which separated two major palaeocontinents, Gondwana and Laurussia (Laurentia-Baltica-Avalonia). One of the principal oceans of the Palaeozoic, its sutures today stretch from Mexico to Turkey and its closure resulted ...
.


Laurussia

By the early Devonian, the continent Laurussia (also known as
Euramerica Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
) was fully formed through the collision of the continents Laurentia (modern day North America) and
Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, is mo ...
(modern day northern and eastern Europe). The tectonic effects of this collision continued into the Devonian, producing a string of mountain ranges along the southeastern coast of the continent. In present-day eastern North America, the Acadian Orogeny continued to raise the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...
. Further east, the collision also extended the rise of the
Caledonian Mountains The Caledonian orogeny was a mountain-building era recorded in the northern parts of the British Isles, the Scandinavian Mountains, Svalbard, eastern Greenland and parts of north-central Europe. The Caledonian orogeny encompasses events that occ ...
of Great Britain and Scandinavia. As the Caledonian Orogeny wound down in the later part of the period,
orogenic collapse In geology, orogenic collapse is the thinning and lateral spread of thickened crust. It is a broad term referring to processes which distribute material from regions of high gravitational potential energy to regions of low gravitational potential ...
facilitated a cluster of granite intrusions in Scotland. Most of Laurussia was located south of the equator, but in the Devonian it moved northwards and began to rotate counterclockwise towards its modern position. While the most northern parts of the continent (such as Greenland and Ellesmere Island) established tropical conditions, most of the continent was located within the natural dry zone along the Tropic of Capricorn, which (as nowadays) is a result of the convergence of two great air-masses, the
Hadley cell The Hadley cell, named after George Hadley, is a global-scale tropical atmospheric circulation that features air rising near the equator, flowing poleward at a height of 10 to 15 kilometers above the earth's surface, descending in the subtropics, ...
and the Ferrel cell. In these near-deserts, the
Old Red Sandstone The Old Red Sandstone is an assemblage of rocks in the North Atlantic region largely of Devonian age. It extends in the east across Great Britain, Ireland and Norway, and in the west along the northeastern seaboard of North America. It also exte ...
sedimentary beds formed, made red by the oxidised iron ( hematite) characteristic of drought conditions. The abundance of red sandstone on continental land also lends Laurussia the name "the Old Red Continent". For much of the Devonian, the majority of western Laurussia (North America) was covered by subtropical
inland seas An inland sea (also known as an epeiric sea or an epicontinental sea) is a continental body of water which is very large and is either completely surrounded by dry land or connected to an ocean by a river, strait, or "arm of the sea". An inland se ...
which hosted a diverse ecosystem of reefs and marine life. Devonian marine deposits are particularly prevalent in the midwestern and northeastern United States. Devonian reefs also extended along the southeast edge of Laurussia, a coastline now corresponding to southern England, Belgium, and other mid-latitude areas of Europe. In the Early and Middle Devonian, the west coast of Laurussia was a passive margin with broad coastal waters, deep silty embayments, river deltas and estuaries, found today in
Idaho Idaho ( ) is a state in the Pacific Northwest region of the Western United States. To the north, it shares a small portion of the Canada–United States border with the province of British Columbia. It borders the states of Montana and Wyomi ...
and Nevada. In the Late Devonian, an approaching volcanic island arc reached the steep slope of the continental shelf and began to uplift deep water deposits. This minor collision sparked the start of a mountain-building episode called the
Antler orogeny The Antler orogeny was a tectonic event that began in the early Late Devonian with widespread effects continuing into the Mississippian and early Pennsylvanian. Most of the evidence for this event is in Nevada but the limits of its reach are unk ...
, which extended into the Carboniferous. Mountain building could also be found in the far northeastern extent of the continent, as minor tropical island arcs and detached Baltic terranes re-join the continent. Deformed remnants of these mountains can still be found on Ellesmere Island and Svalbard. Many of the Devonian collisions in Laurussia produce both mountain chains and
foreland basin A foreland basin is a structural basin that develops adjacent and parallel to a mountain belt. Foreland basins form because the immense mass created by crustal thickening associated with the evolution of a mountain belt causes the lithospher ...
s, which are frequently fossiliferous.


Gondwana

Gondwana was by far the largest continent on the planet. It was completely south of the equator, although the northeastern sector (now Australia) did reach tropical latitudes. The southwestern sector (now South America) was located to the far south, with Brazil situated near the South Pole. The northwestern edge of Gondwana was an active margin for much of the Devonian, and saw the
accretion Accretion may refer to: Science * Accretion (astrophysics), the formation of planets and other bodies by collection of material through gravity * Accretion (meteorology), the process by which water vapor in clouds forms water droplets around nucl ...
of many smaller land masses and island arcs. These include
Chilenia Chilenia was an ancient microcontinent or terrane whose history affected many of the older rocks of central Chile and western Argentina. It was once separated by oceanic crust from the Cuyania terrane to which it accreted at ~420-390 Ma when Cuya ...
,
Cuyania The Precordillera Terrane or Cuyania was an ancient microcontinent or terrane whose history affected many of the older rocks of Cuyo in Argentina. It was separated by oceanic crust from the Chilenia terrane which accreted into it at ~420-390 Ma ...
, and Chaitenia, which now form much of
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
and Patagonia. These collisions were associated with
volcanic activity Volcanism, vulcanism or volcanicity is the phenomenon of eruption of molten rock (magma) onto the surface of the Earth or a solid-surface planet or moon, where lava, pyroclastics, and volcanic gases erupt through a break in the surface called a ...
and plutons, but by the Late Devonian the tectonic situation had relaxed and much of South America was covered by shallow seas. These south polar seas hosted a distinctive brachiopod fauna, the Malvinokaffric Realm, which extended eastward to marginal areas now equivalent to South Africa and Antarctica. Malvinokaffric faunas even managed to approach the South Pole via a tongue of Panthalassa which extended into the Paraná Basin. The northern rim of Gondwana was mostly a passive margin, hosting extensive marine deposits in areas such as northwest Africa and Tibet. The eastern margin, though warmer than the west, was equally active. Numerous mountain building events and granite and kimberlite intrusions affected areas equivalent to modern day eastern Australia, Tasmania, and Antarctica.


Asian terranes

Several island microcontinents (which would later coalesce into modern day Asia) stretched over a low-latitude archipelago to the north of Gondwana. They were separated from the southern continent by an oceanic basin: the Paleo-Tethys. Although the western Paleo-Tethys Ocean had existed since the Cambrian, the eastern part only began to rift apart as late as the Silurian. This process accelerated in the Devonian. The eastern branch of the Paleo-Tethys was fully opened when
South China South China () is a geographical and cultural region that covers the southernmost part of China. Its precise meaning varies with context. A notable feature of South China in comparison to the rest of China is that most of its citizens are not n ...
and Annamia (a terrane equivalent to most of Indochina), together as a unified continent, detached from the northeastern sector of Gondwana. Nevertheless, they remained close enough to Gondwana that their Devonian fossils were more closely related to Australian species than to north Asian species. Other Asian terranes remained attached to Gondwana, including Sibumasu (western Indochina), Tibet, and the rest of the
Cimmerian The Cimmerians ( Akkadian: , romanized: ; Hebrew: , romanized: ; Ancient Greek: , romanized: ; Latin: ) were an ancient Eastern Iranian equestrian nomadic people originating in the Caspian steppe, part of whom subsequently migrated into West ...
blocks. While the South China-Annamia continent was the newest addition to the Asian microcontinents, it was not the first. North China and the Tarim Block (now northwesternmost China) were located westward and continued to drift northwards, powering over older oceanic crust in the process. Further west was a small ocean (the Turkestan Ocean), followed by the larger microcontinents of
Kazakhstania Kazakhstania ( kk, Qazaqstaniya), the Kazakh terranes, or the Kazakhstan Block, is a geological region in Central Asia which consists of the area roughly centered on Lake Balkhash, north and east of the Aral Sea, south of the Siberian craton and ...
, Siberia, and
Amuria Amuria is a town in the Eastern Region of Uganda. It is the chief municipal, administrative, and commercial center of Amuria District, in the Teso sub-region. Location Amuria is located approximately , by road, north of Soroti, the largest c ...
. Kazakhstania was a volcanically active region during the Devonian, as it continued to assimilate smaller island arcs. Siberia was located just north of the equator as the largest landmass in the Northern Hemisphere. At the beginning of the Devonian, Siberia was inverted (upside down) relative to its modern orientation. Later in the period it moved northwards and began to twist clockwise, though it was not near its modern location. Siberia approached the eastern edge of Laurussia as the Devonian progressed, but it was still separated by a seaway, the Ural Ocean. Although Siberia's margins were generally tectonically stable and ecologically productive, rifting and deep mantle plumes impacted the continent with flood basalts during the Late Devonian. The
Altai-Sayan region The Altai-Sayan region is an area of Inner Asia proximate to the Altai Mountains and the Sayan Mountains, near to where Russia, China, Mongolia and Kazakhstan come together. This region is one of the world centers of temperate plant diversity. Its ...
was shaken by volcanism in the Early and Middle Devonian, while Late Devonian magmatism was magnified further to produce the
Vilyuy Traps The Vilyuy ( rus, Вилю́й, p=vʲɪˈlʲʉj; sah, Бүлүү, ''Bülüü'', ) is a river in Russia, the longest tributary of the Lena. About long, it flows mostly within the Sakha Republic. Its basin covers about . History The river is fir ...
, flood basalts which may have contributed to the Late Devonian Mass Extinction. The last major round of volcanism, the Yakutsk Large Igneous Province, continued into the Carboniferous to produce extensive kimberlite deposits. Similar volcanic activity also affected the nearby microcontinent of Amuria (now Manchuria, Mongolia and their vicinities). Though certainly close to Siberia in the Devonian, the precise location of Amuria is uncertain due to contradictory paleomagnetic data.


Closure of the Rheic Ocean

The Rheic Ocean, which separated Laurussia from Gondwana, was wide at the start of the Devonian, having formed after the drift of
Avalonia Avalonia was a microcontinent in the Paleozoic era. Crustal fragments of this former microcontinent underlie south-west Great Britain, southern Ireland, and the eastern coast of North America. It is the source of many of the older rocks of Wester ...
away from Gondwana. It steadily shrunk as the period continued, as the two major continents approached near the equator in the early stages of the assembly of
Pangaea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
. The closure of the Rheic Ocean began in the Devonian and continued into the Carboniferous. As the ocean narrowed, endemic marine faunas of Gondwana and Laurussia combined into a single tropical fauna. The history of the western Rheic Ocean is a subject of debate, but there is good evidence that Rheic oceanic crust experienced intense
subduction Subduction is a geological process in which the oceanic lithosphere is recycled into the Earth's mantle at convergent boundaries. Where the oceanic lithosphere of a tectonic plate converges with the less dense lithosphere of a second plate, ...
and metamorphism under Mexico and Central America. The closure of the eastern part of the Rheic Ocean is associated with the assemblage of central and southern Europe. In the early Paleozoic, much of Europe was still attached to Gondwana, including the terranes of Iberia,
Armorica Armorica or Aremorica (Gaulish: ; br, Arvorig, ) is the name given in ancient times to the part of Gaul between the Seine and the Loire that includes the Brittany Peninsula, extending inland to an indeterminate point and down the Atlantic Coast. ...
(France), Palaeo-Adria (the western Mediterranean area),
Bohemia Bohemia ( ; cs, Čechy ; ; hsb, Čěska; szl, Czechy) is the westernmost and largest historical region of the Czech Republic. Bohemia can also refer to a wider area consisting of the historical Lands of the Bohemian Crown ruled by the Bohe ...
,
Franconia Franconia (german: Franken, ; Franconian dialect: ''Franggn'' ; bar, Frankn) is a region of Germany, characterised by its culture and Franconian dialect (German: ''Fränkisch''). The three administrative regions of Lower, Middle and Upper ...
, and Saxothuringia. These continental blocks, collectively known as the Armorican Terrane Assemblage, split away from Gondwana in the Silurian and drifted towards Laurussia through the Devonian. Their collision with Laurussia leads to the beginning of the Variscan Orogeny, a major mountain-building event which would escalate further in the Late Paleozoic. Franconia and Saxothuringia collided with Laurussia near the end of the Early Devonian, pinching out the easternmost Rheic Ocean. The rest of the Armorican terranes followed, and by the end of the Devonian they were fully connected with Laurussia. This sequence of rifting and collision events led to the successive creation and destruction of several small seaways, including the Rheno-Hercynian, Saxo-Thuringian, and Galicia-Moldanubian oceans. Their sediments were eventually compressed and completely buried as Gondwana fully collided with Laurussia in the Carboniferous.


Life


Marine biota

Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
, diverse and abundant
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, wh ...
s, the enigmatic
hederellid Hederellids are extinct colonial animals with calcitic tubular branching exoskeletons. They range from the Silurian to the Permian and were most common in the Devonian period. They are more properly known as "hederelloids" because they were origin ...
s,
microconchids The order Microconchida is a group of small, spirally-coiled, encrusting fossil "worm" tubes from the class Tentaculita found from the Upper Ordovician to the Middle Jurassic (Bathonian) around the world. They have lamellar calcitic shells ...
and corals. Lily-like
crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
s (animals, their resemblance to flowers notwithstanding) were abundant, and trilobites were still fairly common.
Bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biv ...
became commonplace in deep water and outer shelf environments. The first ammonites also appeared during or slightly before the early Devonian Period around 400  Mya. Bactritoids make their first appearance in the Early Devonian as well; their radiation, along with that of ammonoids, has been attributed by some authors to increased environmental stress resulting from decreasing oxygen levels in the deeper parts of the water column. Among vertebrates, jawless armored fish (
ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and ...
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 Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
) and bony fishes ( Osteichthyes) also become diverse and played a large role within the Devonian seas. The first abundant genus of shark, ''
Cladoselache ''Cladoselache'' is an extinct genus of shark-like chondrichthyan (cartilaginous fish) from the Late Devonian (Famennian) of North America. It was similar in body shape to modern lamnid sharks (such as mako sharks and the great white shark), but ...
'', 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 Devonian saw significant expansion in the diversity of nektonic marine life driven by the abundance of planktonic microorganisms in the free water column as well as high ecological competition in benthic habitats, which were extremely saturated; this diversification has been labeled the Devonian Nekton Revolution by many researchers. However, other researchers have questioned whether this revolution existed at all; a 2018 study found that although the proportion of biodiversity constituted by nekton increased across the boundary between the Silurian and Devonian, it decreased across the span of the Devonian, particularly during the Pragian, and that the overall diversity of nektonic taxa did not increase significantly during the Devonian compared to during other geologic periods, and was in fact higher during the intervals spanning from the Wenlock to the Lochkovian and from the Carboniferous to the Permian. The study's authors instead attribute the increased overall diversity of nekton in the Devonian to a broader, gradual trend of nektonic diversification across the entire Palaeozoic.


Reefs

A now-dry barrier reef, located in present-day Kimberley Basin of northwest Australia, once extended , fringing a Devonian continent. Reefs are generally built by various
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate g ...
-secreting organisms that can erect wave-resistant structures near sea level. Although modern reefs are constructed mainly by corals and calcareous
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
, Devonian reefs were either microbial reefs built up mostly by
autotrophic An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Work ...
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 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, trigonotarbids and
myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
(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 The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
). By far the largest land organism at the beginning of this period was the enigmatic ''
Prototaxites ''Prototaxites'' is a genus of terrestrial fossil fungi dating from the Middle Ordovician until the Late Devonian periods, approximately . ''Prototaxites'' formed small to large trunk-like structures up to wide, reaching in length, made up of ...
'', 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
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsid ...
s 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 ''Piscichnus'' is an ichnogenus of trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or anima ...
'') has been advanced.


The greening of land

Many
Early Devonian plants Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...
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 ''Cooksonia'' is an extinct group of primitive land plants, treated as a genus, although probably not monophyletic. The earliest ''Cooksonia'' date from the middle of the Silurian (the Wenlock epoch); the group continued to be an important comp ...
'' consisted of leafless, dichotomous axes and terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall. Fossils of '' 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:
lycophytes The lycophytes, when broadly circumscribed, are a vascular plant (tracheophyte) subgroup of the kingdom Plantae. They are sometimes placed in a division Lycopodiophyta or Lycophyta or in a subdivision Lycopodiophytina. They are one of the oldest ...
, horsetails, ferns, and
progymnosperm The progymnosperms are an extinct group of woody, spore-bearing plants that is presumed to have evolved from the trimerophytes, and eventually gave rise to the gymnosperms, ancestral to acrogymnosperms and angiosperms (flowering plants). They h ...
s 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 cladoxylopsids and progymnosperm ''
Archaeopteris ''Archaeopteris'' is an extinct genus of progymnosperm tree with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous (), the oldest fossils being 385 million years old, and ...
''. These
tracheophyte Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They al ...
s 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 many plant groups and growth forms has been referred to as the Devonian Explosion or the Silurian-Devonian Terrestrial Revolution. The 'greening' of the continents acted as a
carbon sink A carbon sink is anything, natural or otherwise, that accumulates and stores some carbon-containing chemical compound for an indefinite period and thereby removes carbon dioxide () from the atmosphere. Globally, the two most important carbon ...
, and 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 The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major ex ...
).


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 characterized a recognizably 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 terrestrial living. By the end of the Devonian, arthropods were solidly established on the land.


Gallery

Image:D_Terrelli.png, ''
Dunkleosteus ''Dunkleosteus'' is an extinct genus of large armored, jawed fishes that existed during the Late Devonian period, about 382–358 million years ago. It consists of ten species, some of which are among the largest placoderms to have ever lived: ...
'', 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 ''Cladoselache'' is an extinct genus of shark-like chondrichthyan (cartilaginous fish) from the Late Devonian (Famennian) of North America. It was similar in body shape to modern lamnid sharks (such as mako sharks and the great white shark), but ...
'', several
lobe-finned fishes Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...
, including '' Eusthenopteron'' that was an early marine tetrapod, and the placoderm ''
Bothriolepis ''Bothriolepis'' (from el, βόθρος , 'trench' and el, λεπίς 'scale') was a widespread, abundant and diverse genus of antiarch placoderms that lived during the Middle to Late Devonian period of the Paleozoic Era. Historically, ''Bot ...
'' in a painting from 1905 Image:Melocrinites nodosus spinosus.jpg, ''Melocrinites nodosus spinosus'', a spiny, stalked
crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
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 ''Aulopora'' is an extinct genus of tabulate coral characterized by a bifurcated budding pattern and conical corallite A corallite is the skeletal cup, formed by an individual stony coral polyp, in which the polyp sits and into which it can r ...
'' from the Middle Devonian of Ohio – view of colony encrusting a
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, wh ...
valve Image:Tropidoleptus carinatus.jpg, ''Tropidoleptus carinatus'', an orthid brachiopod from the Middle Devonian of New York Image:Pleurodictyum americanum Kashong.jpg, '' 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 The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major ex ...
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
acritarch Acritarchs are organic microfossils, known from approximately 1800 million years ago to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity refle ...
s, and the world saw the disappearance of an estimated 96% of vertebrates like
conodont Conodonts (Greek ''kōnos'', " cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, whic ...
s and
bony fishes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilage ...
, 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 The Alamo bolide impact occurred 377–378 million years ago, when one or more hypervelocity objects from space slammed into shallow marine waters at a site that is now the Devonian Guilmette Formation of the Worthington Mountains and Schell Cre ...
), little evidence supports the existence of a large enough Devonian crater.


See also

* . 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 * * * * * {{Authority control Geological periods .