Devonian

Devonian

Overview
The Devonian is a geologic period and system of the Paleozoic
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...

 Era spanning from the end of the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...

 Period, about 416.0 ± 2.8 Mya (million years ago), to the beginning of the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

 Period, about 359.2 ± 2.5 Mya . It is named after Devon
Devon
Devon is a large county in southwestern England. The county is sometimes referred to as Devonshire, although the term is rarely used inside the county itself as the county has never been officially "shired", it often indicates a traditional or historical context.The county shares borders with...

, England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

, where rocks from this period were first studied.

The Devonian period experienced the first significant adaptive radiation
Adaptive radiation
In evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...

 of terrestrial life. The first seed-bearing plants
Spermatophyte
The spermatophytes comprise those plants that produce seeds. They are a subset of the embryophytes or land plants...

 spread across dry land.
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Encyclopedia
The Devonian is a geologic period and system of the Paleozoic
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...

 Era spanning from the end of the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...

 Period, about 416.0 ± 2.8 Mya (million years ago), to the beginning of the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

 Period, about 359.2 ± 2.5 Mya . It is named after Devon
Devon
Devon is a large county in southwestern England. The county is sometimes referred to as Devonshire, although the term is rarely used inside the county itself as the county has never been officially "shired", it often indicates a traditional or historical context.The county shares borders with...

, England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

, where rocks from this period were first studied.

The Devonian period experienced the first significant adaptive radiation
Adaptive radiation
In evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...

 of terrestrial life. The first seed-bearing plants
Spermatophyte
The spermatophytes comprise those plants that produce seeds. They are a subset of the embryophytes or land plants...

 spread across dry land. Since large vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

 terrestrial herbivores had not yet appeared, they formed extensive forest
Forest
A forest, also referred to as a wood or the woods, is an area with a high density of trees. As with cities, depending where you are in the world, what is considered a forest may vary significantly in size and have various classification according to how and what of the forest is composed...

s which covered the continents. The first ray finned
Actinopterygii
The Actinopterygii or ray-finned fishes constitute a class or sub-class of the bony fishes.The ray-finned fishes are so called because they possess lepidotrichia or "fin rays", their fins being webs of skin supported by bony or horny spines , as opposed to the fleshy, lobed fins that characterize...

 and lobe-finned
Sarcopterygii
The Sarcopterygii or lobe-finned fishes – sometimes considered synonymous with Crossopterygii constitute a clade of the bony fishes, though a strict classification would include the terrestrial vertebrates...

 bony fish also appeared. The pectoral and pelvic fins of lobe-finned fish evolved into legs as they started to walk on land
Landform
A landform or physical feature in the earth sciences and geology sub-fields, comprises a geomorphological unit, and is largely defined by its surface form and location in the landscape, as part of the terrain, and as such, is typically an element of topography...

 as tetrapod
Tetrapod
Tetrapods are vertebrate animals having four limbs. Amphibians, reptiles, birds and mammals are all tetrapods; even snakes and other limbless reptiles and amphibians are tetrapods by descent. The earliest tetrapods evolved from the lobe-finned fishes in the Devonian...

s around 397 million years ago. They would become the ancestors to early amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

s. Various terrestrial arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s also became well-established.

In the ocean
Ocean
An ocean is a major body of saline water, and a principal component of the hydrosphere. Approximately 71% of the Earth's surface is covered by ocean, a continuous body of water that is customarily divided into several principal oceans and smaller seas.More than half of this area is over 3,000...

s, primitive shark
Shark
Sharks are a type of fish with a full cartilaginous skeleton and a highly streamlined body. The earliest known sharks date from more than 420 million years ago....

s became more numerous than in the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...

 and the late Ordovician. The first ammonite
Ammonite
Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct subclass within the Molluscan class Cephalopoda which are more closely related to living coleoids Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct...

 mollusks appeared. Trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...

s, the mollusc-like brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...

s, as well as great coral reef
Coral reef
Coral reefs are underwater structures made from calcium carbonate secreted by corals. Coral reefs are colonies of tiny living animals found in marine waters that contain few nutrients. Most coral reefs are built from stony corals, which in turn consist of polyps that cluster in groups. The polyps...

s were still common. The Late Devonian extinction
Late Devonian extinction
The Late Devonian extinction was one of five major extinction events in the history of the Earth's biota. A major extinction, the Kellwasser Event, occurred at the boundary that marks the beginning of the last phase of the Devonian period, the Famennian faunal stage, , about 374 million years ago...

 severely affected marine life.

The paleogeography was dominated by the supercontinent
Supercontinent
In geology, a supercontinent is a landmass comprising more than one continental core, or craton. The assembly of cratons and accreted terranes that form Eurasia qualifies as a supercontinent today.-History:...

 of Gondwana
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...

 to the south, the continent
Continent
A continent is one of several very large landmasses on Earth. They are generally identified by convention rather than any strict criteria, with seven regions commonly regarded as continents—they are : Asia, Africa, North America, South America, Antarctica, Europe, and Australia.Plate tectonics is...

 of Siberia
Siberia (continent)
Siberia is the craton located in the heart of the region of Siberia. Siberia or "Angaraland" is today the Central Siberian Plateau...

 to the north, and the early formation of the small supercontinent of Euramerica
Euramerica
Euramerica was a minor supercontinent created in the Devonian as the result of a collision between the Laurentian, Baltica, and Avalonia cratons .300 million years ago in the Late Carboniferous tropical rainforests lay over the equator of Euramerica...

 in between.

History


The period is named after Devon
Devon
Devon is a large county in southwestern England. The county is sometimes referred to as Devonshire, although the term is rarely used inside the county itself as the county has never been officially "shired", it often indicates a traditional or historical context.The county shares borders with...

, a county in southwestern England, where Devonian outcrops are common. While the rock bed
Stratum
In geology and related fields, a stratum is a layer of sedimentary rock or soil with internally consistent characteristics that distinguish it from other layers...

s that define the start and end of the period are well identified, the exact dates are uncertain. According to the International Commission on Stratigraphy
International Commission on Stratigraphy
The International Commission on Stratigraphy , sometimes referred to by the unofficial "International Stratigraphic Commission" is a daughter or major subcommittee grade scientific daughter organization that concerns itself with stratigraphy, geological, and geochronological matters on a global...

 (Ogg, 2004), the Devonian extends from the end of the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...

 Period 416.0 ± 2.8 Mya, to the beginning of the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

 Period 359.9 ± 2.5 Mya (in North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

, the beginning of the Mississippian subperiod of the Carboniferous) (ICS 2004).

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
Old Red Sandstone
The Old Red Sandstone is a British rock formation of considerable importance to early paleontology. For convenience the short version of the term, 'ORS' is often used in literature on the subject.-Sedimentology:...

 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
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...

 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
Stratum
In geology and related fields, a stratum is a layer of sedimentary rock or soil with internally consistent characteristics that distinguish it from other layers...

 of western Europe
Western Europe
Western Europe is a loose term for the collection of countries in the western most region of the European continents, though this definition is context-dependent and carries cultural and political connotations. One definition describes Western Europe as a geographic entity—the region lying in the...

 and eastern North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

, which at the time straddled the Equator
Equator
An equator is the intersection of a sphere's surface with the plane perpendicular to the sphere's axis of rotation and containing the sphere's center of mass....

 as part of the supercontinent of Euramerica where fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

 signatures of widespread reefs indicate tropical climate
Climate
Climate encompasses the statistics of temperature, humidity, atmospheric pressure, wind, rainfall, atmospheric particle count and other meteorological elemental measurements in a given region over long periods...

s that were warm and moderately humid but in fact the climate in the Devonian differed greatly between epoch
Geologic time scale
The geologic time scale provides a system of chronologic measurement relating stratigraphy to time that is used by geologists, paleontologists and other earth scientists to describe the timing and relationships between events that have occurred during the history of the Earth...

s 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
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

 had a warm temperate climate. In the Late Devonian, by contrast, arid conditions were less prevalent across the world and temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 climates were more common.

Subdivisions



The Devonian Period is formally broken into Early, Middle, and Late subdivisions. The rocks corresponding to these epoch
Series (stratigraphy)
Series are subdivisions of rock layers made based on the age of the rock and corresponding to the dating system unit called an epoch, both being formally defined international conventions of the geological timescale. A series is therefore a sequence of rock depositions defining a...

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

The Early Devonian lasts from and begins with the Lochkovian stage, which lasts until the Pragian. This spans from , and is followed by the Emsian, which lasts until the Middle Devonian begins, .
The Middle Devonian comprises two subdivisions, the Eifelian giving way to the Givetian .
During this time the armoured jawless ostracoderm
Ostracoderm
Ostracoderms are any of several groups of extinct, primitive, jawless fishes that were covered in an armor of bony plates. They belong to the taxon Ostracodermi, and their fossils are found in the Ordovician and Devonian Period strata of North America and Europe...

 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, , 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, .

Climate


The Devonian was a relatively warm period, and probably lacked any glaciers. Reconstruction of tropical sea surface temperature from conodont
Conodont
Conodonts are extinct chordates resembling eels, classified in the class Conodonta. For many years, they were known only from tooth-like microfossils now called conodont elements, found in isolation. Knowledge about soft tissues remains relatively sparse to this day...

 apatite
Apatite
Apatite is a group of phosphate minerals, usually referring to hydroxylapatite, fluorapatite, chlorapatite and bromapatite, named for high concentrations of OH−, F−, Cl− or Br− ions, respectively, in the crystal...

 implies an average value of 30 °C (86 °F) in the Early Devonian. 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-change. 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 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
Plate tectonics
Plate tectonics is a scientific theory that describes the large scale motions of Earth's lithosphere...

 activity, as Euramerica
Euramerica
Euramerica was a minor supercontinent created in the Devonian as the result of a collision between the Laurentian, Baltica, and Avalonia cratons .300 million years ago in the Late Carboniferous tropical rainforests lay over the equator of Euramerica...

 and Gondwana
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...

 drew closer together.

The continent Euramerica
Euramerica
Euramerica was a minor supercontinent created in the Devonian as the result of a collision between the Laurentian, Baltica, and Avalonia cratons .300 million years ago in the Late Carboniferous tropical rainforests lay over the equator of Euramerica...

 (or Laurussia) was created in the early Devonian by the collision of Laurentia
Laurentia
Laurentia is a large area of continental craton, which forms the ancient geological core of the North American continent...

 and Baltica
Baltica
Baltica is a name applied by geologists to a late-Proterozoic, early-Palaeozoic continent that now includes the East European craton of northwestern Eurasia. Baltica was created as an entity not earlier than 1.8 billion years ago. Before this time, the three segments/continents that now comprise...

, which rotated into the natural dry zone along the Tropic of Capricorn
Tropic of Capricorn
The Tropic of Capricorn, or Southern tropic, marks the most southerly latitude on the Earth at which the Sun can be directly overhead. This event occurs at the December solstice, when the southern hemisphere is tilted towards the Sun to its maximum extent.Tropic of Capricorn is one of the five...

, which is formed as much in Paleozoic times as nowadays by the convergence of two great air-masses, the Hadley cell
Hadley cell
The Hadley cell, named after George Hadley, is a circulation pattern that dominates the tropical atmosphere, with rising motion near the equator, poleward flow 10–15 kilometers above the surface, descending motion in the subtropics, and equatorward flow near the surface...

 and the Ferrel cell. In these near-deserts, the Old Red Sandstone
Old Red Sandstone
The Old Red Sandstone is a British rock formation of considerable importance to early paleontology. For convenience the short version of the term, 'ORS' is often used in literature on the subject.-Sedimentology:...

 sedimentary beds formed, made red by the oxidized iron (hematite
Hematite
Hematite, also spelled as haematite, is the mineral form of iron oxide , one of several iron oxides. Hematite crystallizes in the rhombohedral system, and it has the same crystal structure as ilmenite and corundum...

) characteristic of drought conditions.

Near the equator
Equator
An equator is the intersection of a sphere's surface with the plane perpendicular to the sphere's axis of rotation and containing the sphere's center of mass....

, the plate
Tectonic Plates
Tectonic Plates is a 1992 independent Canadian film directed by Peter Mettler. Mettler also wrote the screenplay based on the play by Robert Lepage. The film stars Marie Gignac, Céline Bonnier and Robert Lepage.-Plot summary:...

 of Euramerica and Gondwana were starting to meet, beginning the early stages of assembling Pangaea
Pangaea
Pangaea, Pangæa, or Pangea is hypothesized as a supercontinent that existed during the Paleozoic and Mesozoic eras about 250 million years ago, before the component continents were separated into their current configuration....

. This activity further raised the northern Appalachian Mountains
Appalachian Mountains
The Appalachian Mountains #Whether the stressed vowel is or ,#Whether the "ch" is pronounced as a fricative or an affricate , and#Whether the final vowel is the monophthong or the diphthong .), often called the Appalachians, are a system of mountains in eastern North America. The Appalachians...

 and formed the Caledonian Mountains
Caledonian orogeny
The Caledonian orogeny is 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 occurred from the Ordovician to Early Devonian, roughly...

 in Great Britain
Great Britain
Great Britain or Britain is an island situated to the northwest of Continental Europe. It is the ninth largest island in the world, and the largest European island, as well as the largest of the British Isles...

 and Scandinavia
Scandinavia
Scandinavia is a cultural, historical and ethno-linguistic region in northern Europe that includes the three kingdoms of Denmark, Norway and Sweden, characterized by their common ethno-cultural heritage and language. Modern Norway and Sweden proper are situated on the Scandinavian Peninsula,...

.

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
Idaho
Idaho is a state in the Rocky Mountain area of the United States. The state's largest city and capital is Boise. Residents are called "Idahoans". Idaho was admitted to the Union on July 3, 1890, as the 43rd state....

 and Nevada
Nevada
Nevada is a state in the western, mountain west, and southwestern regions of the United States. With an area of and a population of about 2.7 million, it is the 7th-largest and 35th-most populous state. Over two-thirds of Nevada's people live in the Las Vegas metropolitan area, which contains its...

; an approaching volcanic island arc
Island arc
An island arc is a type of archipelago composed of a chain of volcanoes which alignment is arc-shaped, and which are situated parallel and close to a boundary between two converging tectonic plates....

 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
Antler orogeny
The Antler orogeny is a mountain-building episode that is named for Antler Peak, at Battle Mountain, Nevada. The orogeny extensively deformed Paleozoic rocks of the Great Basin in Nevada and western Utah during Late Devonian and Early Mississippian time...

.

Sea levels were high worldwide, and much of the land lay submerged under shallow seas, where tropical reef
Reef
In nautical terminology, a reef is a rock, sandbar, or other feature lying beneath the surface of the water ....

 organisms lived. The deep, enormous Panthalassa
Panthalassa
Panthalassa , also known as the Panthalassic Ocean, was the vast global ocean that surrounded the supercontinent Pangaea, during the late Paleozoic and the early Mesozoic years. It included the Pacific Ocean to the west and north and the Tethys Ocean to the southeast...

 (the "universal ocean") covered the rest of the planet
Planet
A planet is a celestial body orbiting a star or stellar remnant that is massive enough to be rounded by its own gravity, is not massive enough to cause thermonuclear fusion, and has cleared its neighbouring region of planetesimals.The term planet is ancient, with ties to history, science,...

. Other minor oceans were Paleo-Tethys, Proto-Tethys, Rheic Ocean
Rheic Ocean
The Rheic Ocean was a Paleozoic ocean between the large continent Gondwana to the south and the microcontinents Avalonia and others to the north...

, and Ural Ocean
Ural Ocean
The Ural Ocean was a small, ancient ocean that was situated between Siberia and Baltica. The ocean formed in the Late Ordovician epoch, when large islands from Siberia collided with Baltica, which was now part of a minor supercontinent of Euramerica. The islands also caused Ural Ocean's precursor,...

 (which was closed during the collision with Siberia
Siberia (continent)
Siberia is the craton located in the heart of the region of Siberia. Siberia or "Angaraland" is today the Central Siberian Plateau...

 and Baltica).

Marine biota


Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by bryozoa
Bryozoa
The Bryozoa, also known as Ectoprocta or commonly as moss animals, are a phylum of aquatic invertebrate animals. Typically about long, they are filter feeders that sieve food particles out of the water using a retractable lophophore, a "crown" of tentacles lined with cilia...

, diverse and abundant brachiopods, the enigmatic hederelloids
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 originally defined as a suborder by Bassler , who...

, microconchids and coral
Coral
Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of...

s. Lily-like crinoids were abundant, and trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...

s were still fairly common. Among vertebrates, jaw-less armored fish (ostracoderms) declined in diversity, while the jawed fish (gnathostomes) simultaneously increased in both the sea and fresh water
Fresh Water
Fresh Water is the debut album by Australian rock and blues singer Alison McCallum, released in 1972. Rare for an Australian artist at the time, it came in a gatefold sleeve...

. 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
Chondrichthyes or cartilaginous fishes are jawed fish with paired fins, paired nares, scales, two-chambered hearts, and skeletons made of cartilage rather than bone...

) and bony fishes (Osteichthyes
Osteichthyes
Osteichthyes , also called bony fish, are a taxonomic group of fish that have bony, as opposed to cartilaginous, skeletons. The vast majority of fish are osteichthyes, which is an extremely diverse and abundant group consisting of over 29,000 species...

) also become diverse and played a large role within the Devonian seas. The first abundant genus of shark, Cladoselache
Cladoselache
Cladoselache is a genus of extinct shark. It appeared in the Devonian period.This primitive shark grew to be up to long and roamed the oceans of North America. It is known to have been a fast moving and fairly agile predator due to its streamlined body and deep forked tail...

, 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 ammonite
Ammonite
Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct subclass within the Molluscan class Cephalopoda which are more closely related to living coleoids Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct...

s 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
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, once extended a thousand kilometers, fringing a Devonian continent. Reefs in general are built by various carbonate
Carbonate
In chemistry, a carbonate is a salt of carbonic acid, characterized by the presence of the carbonate ion, . The name may also mean an ester of carbonic acid, an organic compound containing the carbonate group C2....

-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
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

. They were composed of calcareous algae and coral-like stromatoporoids
Stromatoporoidea
Stromatoporoidea is a class of aquatic invertebrates common in the fossil record from the Ordovician through the Cretaceous. They were especially abundant in the Silurian and Devonian. These invertebrates were important reef-formers throughout the Paleozoic and the Late Mesozoic. The group was...

, and tabulate and rugose corals
Rugosa
Disambiguation:The Rugosa Rose is also sometimes just called "Rugosa". For the moon in Star Wars: The Clone Wars, see .The Rugosa, also called the Tetracoralla, are an extinct order of coral that were abundant in Middle Ordovician to Late Permian seas.Solitary rugosans are often referred to...

, in that order of importance.



Terrestrial biota


By the Devonian Period, life was well underway in its colonization of the land. The moss
Moss
Mosses are small, soft plants that are typically 1–10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems...

 forests and bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

l and algal mats of the Silurian
Silurian
The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...

 were joined early in the period by primitive rooted plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s that created the first stable soil
Soil
Soil is a natural body consisting of layers of mineral constituents of variable thicknesses, which differ from the parent materials in their morphological, physical, chemical, and mineralogical characteristics...

s and harbored arthropods like mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...

s, scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...

s and myriapods (although arthropods appeared on land much earlier than in the Early Devonian and the existence of fossils such as Climactichnites
Climactichnites
Climactichnites is an enigmatic, late Cambrian fossil formed on or within sandy tidal flats around .It has been interpreted in many different ways in the past, but is now known to be a trace fossil of a slug-like organism, thought to have moved by crawling on near-shore or on-shore surfaces or...

suggest that land arthropods may have appeared as early as the Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...

 period). 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 greatest land organism was Prototaxites
Prototaxites
The genus Prototaxites describes terrestrial organisms known only from fossils dating from the Silu-Devonian, approximately 420 to 370 million years ago. Prototaxites formed large trunk-like structures up to wide, reaching in height, made up of interwoven tubes just in diameter...

, the fruiting body of an enormous fungus that stood more than 8 meters tall, towering over the low, carpet-like vegetation. By Middle Devonian, shrub-like forests of primitive plants existed: lycophytes, horsetails, fern
Fern
A fern is any one of a group of about 12,000 species of plants belonging to the botanical group known as Pteridophyta. Unlike mosses, they have xylem and phloem . They have stems, leaves, and roots like other vascular plants...

s, and progymnosperms had evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

. Most of these plants had true roots and leaves, and many were quite tall. The earliest known trees, from the genus Wattieza
Wattieza
Wattieza was a genus of prehistoric trees that existed in the mid-Devonian that belong to the cladoxylopsids, close relatives of the modern ferns and horsetails...

, appeared in the Late Devonian around 380 Ma. In the Late Devonian, the tree-like ancestral fern Archaeopteris
Archaeopteris
Archaeopteris is an extinct genus of tree-like plants with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous , and has a global distribution....

and the giant cladoxylopsid
Cladoxylopsid
The cladoxylopsids are a group of plants known only as fossils that are thought to be ancestors of ferns and horsetails.They had a central trunk, from the top of which several lateral branches were attached. Fossils of these plants originate in the Middle Devonian to Early Carboniferous periods ,...

 trees grew with true wood
Wood
Wood is a hard, fibrous tissue found in many trees. It has been used for hundreds of thousands of years for both fuel and as a construction material. It is an organic material, a natural composite of cellulose fibers embedded in a matrix of lignin which resists compression...

. (See also: lignin
Lignin
Lignin or lignen is a complex chemical compound most commonly derived from wood, and an integral part of the secondary cell walls of plants and some algae. The term was introduced in 1819 by de Candolle and is derived from the Latin word lignum, meaning wood...

.) 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
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 sink
Carbon dioxide sink
A carbon sink is a natural or artificial reservoir that accumulates and stores some carbon-containing chemical compound for an indefinite period. The process by which carbon sinks remove carbon dioxide from the atmosphere is known as carbon sequestration...

, and atmospheric
Earth's atmosphere
The atmosphere of Earth is a layer of gases surrounding the planet Earth that is retained by Earth's gravity. The atmosphere protects life on Earth by absorbing ultraviolet solar radiation, warming the surface through heat retention , and reducing temperature extremes between day and night...

 levels of this greenhouse gas
Greenhouse gas
A greenhouse gas is a gas in an atmosphere that absorbs and emits radiation within the thermal infrared range. This process is the fundamental cause of the greenhouse effect. The primary greenhouse gases in the Earth's atmosphere are water vapor, carbon dioxide, methane, nitrous oxide, and ozone...

 may have dropped. This may have cooled the climate and led to a massive extinction event
Extinction event
An extinction event is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation...

. See Late Devonian extinction
Late Devonian extinction
The Late Devonian extinction was one of five major extinction events in the history of the Earth's biota. A major extinction, the Kellwasser Event, occurred at the boundary that marks the beginning of the last phase of the Devonian period, the Famennian faunal stage, , about 374 million years ago...

.

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
Erosion
Erosion is when materials are removed from the surface and changed into something else. It only works by hydraulic actions and transport of solids in the natural environment, and leads to the deposition of these materials elsewhere...

 and sediment deposition. The rapid evolution of a terrestrial ecosystem containing copious animals opened the way for the first vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

s 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, (the Frasnian-Famennian boundary), about 364 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 brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...

s, trilobites, ammonite
Ammonite
Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct subclass within the Molluscan class Cephalopoda which are more closely related to living coleoids Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct...

s, conodont
Conodont
Conodonts are extinct chordates resembling eels, classified in the class Conodonta. For many years, they were known only from tooth-like microfossils now called conodont elements, found in isolation. Knowledge about soft tissues remains relatively sparse to this day...

s, and acritarch
Acritarch
Acritarchs are small organic fossils, present from approximately to the present. Their diversity reflects major ecological events such as the appearance of predation and the Cambrian explosion.-Definition:In general, any small, non-acid soluble Acritarchs are small organic fossils, present from...

s, 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
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...

 paleontologist Digby McLaren
Digby McLaren
Digby Johns McLaren, OC, FRSC was a Canadian geologist and palaeontologist.Born in Carrickfergus, Ireland and educated at Sedbergh School, he received a Bachelor of Arts in geology from the University of Cambridge. During World War II, he fought in the Middle East and Europe with the Royal...

 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
Alamo bolide impact
The Alamo bolide impact occurred 367 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 Creek Range of southeastern Nevada; the event is named for...

), little evidence supports the existence of a Devonian crater large enough.

See also

  • Geologic timescale
  • Phacops rana
    Phacops rana
    Phacops rana is a species of trilobite from the middle Devonian period. Their fossils are found chiefly in the northeastern United States, southwestern Ontario, and in Morocco.-Physiology:...

    : a Devonian trilobite.
  • List of fossil sites (with link directory)

External links