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The Devonian ( ) is a
geologic period The geologic time scale or geological time scale (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochronolo ...
and
system A system is a group of interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its open system (systems theory), environment, is described by its boundaries, str ...
of the
Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
era An era is a span of time. Era or ERA may also refer to: * Era (geology), a subdivision of geologic time * Calendar era Education * Academy of European Law (German: '), an international law school * ERA School, in Melbourne, Australia * E ...
during the
Phanerozoic The Phanerozoic is the current and the latest of the four eon (geology), geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and ...
eon, spanning 60.3 million years from the end of the preceding
Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
period at
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
( Ma), to the beginning of the succeeding
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period at Ma. It is the fourth period of both the Paleozoic and the Phanerozoic. It is named after
Devon Devon ( ; historically also known as Devonshire , ) is a ceremonial county in South West England. It is bordered by the Bristol Channel to the north, Somerset and Dorset to the east, the English Channel to the south, and Cornwall to the west ...
,
South West England South West England, or the South West of England, is one of the nine official regions of England, regions of England in the United Kingdom. Additionally, it is one of four regions that altogether make up Southern England. South West England con ...
, where rocks from this period were first studied. The first significant
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
of life on land occurred during the Devonian, as free- sporing
land plant The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophyt ...
s (
pteridophyte A pteridophyte is a vascular plant (with xylem and phloem) that reproduces by means of spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as " cryptogams", meaning that their means of reproduction is ...
s) began to spread across
dry land Land, also known as dry land, ground, or earth, is the solid terrestrial surface of Earth not submerged by the ocean or another body of water. It makes up 29.2% of Earth's surface and includes all continents and islands. Earth's land surface ...
, forming extensive
coal forest Coal forests were the vast swathes of freshwater swamp and riparian forests that covered much of the lands on Earth's tropical regions during the late Carboniferous ( Pennsylvanian) and Permian periods.Cleal, C. J. & Thomas, B. A. (2005). "Pala ...
s which covered the continents. By the middle of the Devonian, several groups of
vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s had evolved
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
and true
root In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...
s, and by the end of the period the first
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
-bearing plants ( pteridospermatophytes) appeared. This rapid evolution and colonization process, which had begun during the Silurian, is known as the
Silurian-Devonian Terrestrial Revolution The Silurian-Devonian Terrestrial Revolution, also known as the Devonian Plant Explosion (DePE) and the Devonian explosion, was a period of rapid colonization, diversification and radiation of land plants and fungi on dry lands that occurred 428 ...
. The earliest
land animal Terrestrial animals are animals that live predominantly or entirely on land (e.g. cats, chickens, ants, most spiders), as compared with aquatic animals, which live predominantly or entirely in the water (e.g. fish, lobsters, octopuses), and semi ...
s, predominantly
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s such as
myriapod 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. Although molecular evidence and similar fossils suggests a diversifi ...
s,
arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s and hexapods, also became well-established early in this period, after beginning their colonization of land at least from the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
period.
Fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
es, especially
jawed fish Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant taxon, extant vertebrates, including all living bony fishes (bot ...
, reached substantial diversity during this time, leading the Devonian to be called the Age of Fishes. The
armor Armour (Commonwealth English) or armor (American English; see American and British English spelling differences#-our, -or, spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, e ...
ed
placoderm Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...
s began dominating almost every known aquatic environment. In the oceans,
cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
es such as primitive
shark Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s became more numerous than in the Silurian and Late Ordovician.
Tetrapodomorph Tetrapodomorpha (also known as Choanata) is a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced for ...
s, which include the ancestors of all four-
limb Limb may refer to: Science and technology *Limb (anatomy), an appendage of a human or animal *Limb, a large or main branch of a tree *Limb, in astronomy, the curved edge of the apparent disk of a celestial body, e.g. lunar limb *Limb, in botany, t ...
ed vertebrates (i.e.
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s), began diverging from
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...
lobe-finned fish Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
as their more robust and muscled
pectoral Pectoral may refer to: * The chest region and anything relating to it. * Pectoral cross, a cross worn on the chest * a decorative, usually jeweled version of a gorget * Pectoral (Ancient Egypt), a type of jewelry worn in ancient Egypt * Pectora ...
and
pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hi ...
s gradually evolved into
forelimb A forelimb or front limb is one of the paired articulated appendages ( limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used inst ...
s and
hindlimb A hindlimb or back limb is one of the paired articulated appendages ( limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-H ...
s, though they were not fully established for life on land until the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
. The first
ammonite Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
s, a subclass of
cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
s, appeared.
Trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...
s,
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of 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 e ...
s and the great
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in group ...
s were still common during the Devonian. The
Late Devonian extinction The Late Devonian mass extinction, also known as the Kellwasser event, was a mass extinction event which occurred around 372 million years ago, at the boundary between the Frasnian and Famennian ages of the Late Devonian period.Racki, 2005McGh ...
, which started about 375 Ma, severely affected marine life, killing off most of the reef systems, most of the jawless fish, the placoderms, and nearly 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 surviving order of trilobite, dying out in the Permian-Triassic extinction event. Description These typically small trilobites resemble those of ...
. The subsequent
end-Devonian extinction The Hangenberg event, also known as the Hangenberg crisis or end-Devonian extinction, is a mass extinction that occurred at the end of the Famennian stage, the last stage in the Devonian Period (roughly 358.9 ± 0.4 million years ago). It is usu ...
, which occurred at around 359 Ma, further impacted the ecosystems and completed the
extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
of all
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
sponge reef Sponge reefs are reefs produced by sea sponges. All modern sponge reefs are formed by hexactinellid sponges, which have an endoskeleton made of silica spicules and are often referred to as "glass sponges", while historically the non-spicule ...
s and placoderms. Devonian
palaeogeography Palaeogeography (or paleogeography) is the study of historical geography, generally physical landscapes. Palaeogeography can also include the study of human or cultural environments. When the focus is specifically on landforms, the term pale ...
was dominated by the
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
to the south, the small continent of
Siberia Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
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 (Million years ago, Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during ...
to the east. Major tectonic events include the closure of the
Rheic Ocean The Rheic Ocean (; ) was an ocean which separated two major paleocontinents, Gondwana and Laurussia ( Laurentia- Baltica- Avalonia). One of the principal oceans of the Paleozoic, its sutures today stretch from Mexico to Turkey and its closure r ...
, the separation of
South China South China ( zh, s=, p=Huá'nán, j=jyut6 naam4) 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 ...
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; ex ...
. The Devonian experienced several major mountain-building events as Laurussia and Gondwana approached; these include the
Acadian Orogeny The Acadian orogeny is a long-lasting mountain building event which began in the Middle Devonian, reaching a climax in the Late Devonian. It was active for approximately 50 million years, beginning roughly around 375 million years ago (Ma), wi ...
in North America and the beginning of the
Variscan Orogeny The Variscan orogeny, or Hercynian orogeny, was a geologic mountain-building event caused by Late Paleozoic continental collision between Euramerica (Laurussia) and Gondwana to form the supercontinent of Pangaea. Nomenclature The name ''Varis ...
in Europe. These early collisions preceded the formation of the single supercontinent
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 period approximately 335 mi ...
in the Late Paleozoic.


History

The period is named after
Devon Devon ( ; historically also known as Devonshire , ) is a ceremonial county in South West England. It is bordered by the Bristol Channel to the north, Somerset and Dorset to the east, the English Channel to the south, and Cornwall to the west ...
, a county in southwestern England, where a controversial argument in the 1830s over the age and structure of the rocks found throughout the county was resolved by adding the Devonian Period to 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 sin ...
was a lengthy debate between
Roderick Murchison Sir Roderick Impey Murchison, 1st Baronet (19 February 1792 – 22 October 1871) was a Scottish geologist who served as director-general of the British Geological Survey from 1855 until his death in 1871. He is noted for investigating and desc ...
,
Adam Sedgwick Adam Sedgwick FRS (; 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 ...
and
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 ...
over the naming of the period. Murchison and Sedgwick won the debate and named it 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 International Commission on Stratigraphy (ICS), sometimes unofficially referred to as the International Stratigraphic Commission, is a daughter or major subcommittee grade scientific organization that concerns itself with stratigraphy, strati ...
, the Devonian extends from the end of the Silurian Ma, to the beginning of the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
Ma – in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, 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 Old Red Sandstone, abbreviated ORS, 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 eastern seaboard of North America. It ...
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 A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
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 (statistics), bias in which a sample is collected in such a way that some members of the intended statistical population, population have a lower or higher sampling probability than others. It results in a b ...
: most of the early Devonian-age discoveries came from the
strata In geology and related fields, a stratum (: strata) is a layer of Rock (geology), rock or sediment characterized by certain Lithology, lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by v ...
of
Western Europe Western Europe is the western region of Europe. The region's extent varies depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the Western half of the ancient Mediterranean ...
and eastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, which at the time straddled the
Equator The equator is the circle of latitude that divides Earth into the Northern Hemisphere, Northern and Southern Hemisphere, Southern Hemispheres of Earth, hemispheres. It is an imaginary line located at 0 degrees latitude, about in circumferen ...
as part of the supercontinent of Euramerica where
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
signatures of widespread reefs indicate tropical
climate Climate is the long-term weather pattern in a region, 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 meteoro ...
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 South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
had a warm
temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ra ...
. In the Late Devonian, by contrast, arid conditions were less prevalent across the world and
temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ra ...
s were more common.


Subdivisions

The Devonian Period is formally broken into Early, Middle and Late subdivisions. The rocks corresponding to those
epoch 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 b ...
s are referred to as belonging to the Lower, Middle and Upper parts of the Devonian System.


Early Devonian

The
Early Devonian The Early Devonian is the first of three Epoch (geology), epochs comprising the Devonian period, corresponding to the Lower Devonian Series (stratigraphy), series. It lasted from and began with the Lochkovian Stage , which was followed by the Pr ...
lasted from to Ma. It 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 from to Ma, 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 ...
from to Ma and then by the
Emsian The Emsian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.62 ±1.95 million years ago to 393.47 ±0.99 million years ago. It was preceded by the Pragian Stage and followed by the Eifelian Stage. It is named after ...
, which lasted until the Middle Devonian began at Ma. During this time, the first
ammonoids Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
appeared, descending from bactritoid
nautiloids Nautiloids are a group of cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and species rich, with over 2,500 recorded species. Th ...
. Ammonoids during this time period were simple and differed little from their nautiloid counterparts. These ammonoids belong to the order Agoniatitida, which in later epochs evolved to new ammonoid orders, for example
Goniatitida Goniatids, informally goniatites, are ammonoid cephalopods that form the order Goniatitida, derived from the more primitive Agoniatitida during the Middle Devonian some 390 million years ago (around Eifelian stage). Goniatites (goniatitids) su ...
and
Clymeniida Clymeniida is an order of ammonoid cephalopods from the Upper Devonian characterized by having an unusual dorsal siphuncle. They measured about in diameter and are most common in Europe, North Africa, and South China but are known from North ...
. This class of
cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
molluscs would dominate the marine fauna until the beginning of the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
Era.


Middle Devonian

The
Middle Devonian In the geological timescale, the Middle Devonian epoch (from 393.3 ± 1.2 million years ago to 382.7 ± 1.6 million years ago) occurred during the Devonian period, after the end of the Emsian age. The Middle Devonian epoch is subdivided into two ...
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 The Givetian is one of two faunal stages in the Middle Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Eifelian Stage and followed by the Frasnian Stage. It is named after the town of Givet in ...
at Ma. During this time, the jawless agnathan fishes began to decline in diversity in freshwater and marine environments partly due to drastic environmental changes and partly due to the increasing competition, predation, and diversity of jawed fishes. The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop such essential characteristics as well developed lungs and the ability to crawl out of the water and onto the land for short periods of time.


Late Devonian

Finally, the Late Devonian started with the
Frasnian 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 ...
, from to Ma, 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 later of two faunal stages in the Late Devonian epoch. The most recent estimate for its duration is that it lasted from around 371.1 to 359.3 million years ago. An earlier 2012 estimate, still used by the International Commis ...
subdivision, the beginning and end of which are marked with extinction events. This lasted until the end of the Devonian at Ma.


Climate

The Devonian was a relatively warm period, although significant
glacier A glacier (; or ) is a persistent body of dense ice, a form of rock, that is constantly moving downhill under its own weight. A glacier forms where the accumulation of snow exceeds its ablation over many years, often centuries. It acquires ...
s may have existed during the Early and Middle Devonian. 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 Sea surface temperature (or ocean surface temperature) is the ocean temperature, temperature of ocean water close to the surface. The exact meaning of ''surface'' varies in the literature and in practice. It is usually between and below the sea ...
from
conodont Conodonts, are an extinct group of marine jawless vertebrates belonging to the class Conodonta (from Ancient Greek κῶνος (''kōnos''), meaning " cone", and ὀδούς (''odoús''), meaning "tooth"). They are primarily known from their hard ...
apatite Apatite is a group of phosphate minerals, usually hydroxyapatite, fluorapatite and chlorapatite, with high concentrations of Hydroxide, OH−, Fluoride, F− and Chloride, Cl− ion, respectively, in the crystal. The formula of the admixture of ...
implies an average value of in the Early Devonian. Early Devonian mean annual surface temperatures were approximately 16 °C. levels dropped steeply throughout the Devonian Period. The newly evolved forests drew carbon out of the atmosphere, which were then buried into sediments. This may be reflected by a Mid-Devonian cooling of around . The Late Devonian warmed to levels equivalent to the Early Devonian; while there is no corresponding increase in concentrations, continental weathering increases (as predicted by warmer temperatures); further, a range of evidence, such as plant distribution, points to a Late Devonian warming. The climate would have affected the dominant organisms in
reef A reef is a ridge or shoal of rock, coral, or similar relatively stable material lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic component, abiotic (non-living) processes such as deposition (geol ...
s;
microbes A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells. The possible existence of unseen microbial life was suspected from antiquity, with an early attestation in ...
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 Middle Ordovician to the Late Devonian.Stock, C.W. 2001, Stromatoporoidea, 1926–2000: ''Journal of Paleontology'', v. 75, p. 1079–1089. They can be cha ...
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 Paleozoic icehouse The late Paleozoic icehouse, also known as the Late Paleozoic Ice Age (LPIA) and formerly known as the Karoo ice age, was an ice age that began in the Late Devonian and ended in the Late Permian, occurring from 360 to 255 million years ago (Mya), ...
.


Paleogeography

The Devonian world involved many continents and ocean basins of various sizes. The largest continent,
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
, was located entirely within the Southern Hemisphere. It corresponds to modern day
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
,
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
,
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
,
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
, and
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
, as well as minor components of
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
and
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
. The second-largest continent, Laurussia, was northwest of Gondwana, and corresponds to much of modern-day
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
and
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. Various smaller continents, microcontinents, and
terrane In geology, a terrane (; in full, a tectonostratigraphic terrane) is a crust fragment formed on a tectonic plate (or broken off from it) and accreted or " sutured" to crust lying on another plate. The crustal block or fragment preserves its d ...
s 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 Tectonics ( via Latin ) are the processes that result in the structure and properties of the Earth's crust and its evolution through time. The field of ''planetary tectonics'' extends the concept to other planets and moons. These processes ...
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 (Million years ago, Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during ...
and
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
drew closer together. Sea levels were high worldwide, and much of the land lay under shallow seas, where tropical
reef A reef is a ridge or shoal of rock, coral, or similar relatively stable material lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic component, abiotic (non-living) processes such as deposition (geol ...
organisms lived. The enormous "world ocean",
Panthalassa Panthalassa, also known as the Panthalassic Ocean or Panthalassan Ocean (from Greek "all" and "sea"), was the vast superocean that encompassed planet Earth and surrounded the supercontinent Pangaea, the latest in a series of supercontinent ...
, occupied much of the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
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; ex ...
and
Rheic Ocean The Rheic Ocean (; ) was an ocean which separated two major paleocontinents, Gondwana and Laurussia ( Laurentia- Baltica- Avalonia). One of the principal oceans of the Paleozoic, its sutures today stretch from Mexico to Turkey and its closure r ...
.


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 Laurentia or the North American craton is a large continental craton that forms the Geology of North America, ancient geological core of North America. Many times in its past, Laurentia has been a separate continent, as it is now in the form of ...
(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, i ...
(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 The Acadian orogeny is a long-lasting mountain building event which began in the Middle Devonian, reaching a climax in the Late Devonian. It was active for approximately 50 million years, beginning roughly around 375 million years ago (Ma), wi ...
continued to raise the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, are a mountain range in eastern to northeastern North America. The term "Appalachian" refers to several different regions associated with the mountain range, and its surrounding terrain ...
. Further east, the collision also extended the rise of the Caledonian Mountains of
Great Britain Great Britain is an island in the North Atlantic Ocean off the north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
and Scandinavia. As the Caledonian Orogeny wound down in the later part of the period, orogenic collapse 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 and the Ferrel cell. In these near-deserts, the
Old Red Sandstone Old Red Sandstone, abbreviated ORS, 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 eastern seaboard of North America. It ...
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 sea (geology), inland seas which hosted a diverse ecosystem of reefs and marine life. Devonian marine deposits are particularly prevalent in the Midwestern United States, midwestern and Northeastern United States, 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 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, 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 basins, 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 (geology), accretion of many smaller land masses and island arcs. These include Chilenia, Cuyania, and Chaitenia, which now form much of Chile and Tectonic evolution of Patagonia, Patagonia. These collisions were associated with volcanic activity 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 Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
, 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 Ocean, 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 craton, South China and Annamia block, Annamia (a
terrane In geology, a terrane (; in full, a tectonostratigraphic terrane) is a crust fragment formed on a tectonic plate (or broken off from it) and accreted or " sutured" to crust lying on another plate. The crustal block or fragment preserves its d ...
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 Cimmeria (continent), Cimmerian blocks. While the South China-Annamia continent was the newest addition to the Asian microcontinents, it was not the first. North China craton, 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,
Siberia Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
, and Amurian Plate, Amuria. Kazakhstania was a volcanically active region during the Devonian, as it continued to assimilate smaller island arcs. The island arcs of the region, such as the Balkhash-West Junggar Arc, exhibited biological endemism as a consequence of their location. 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 was shaken by volcanism in the Early and Middle Devonian, while Late Devonian magmatism was magnified further to produce the Vilyuy Traps, 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 Paleomagnetism, 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 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 period approximately 335 mi ...
. 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 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, Armorican terrane, Armorica (France), Palaeo-Adria (the western Mediterranean area), Bohemia, Franconia, and Saxothuringian Zone, 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, 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 conodonts, bryozoa, bryozoans, diverse and abundant
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of 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 e ...
s, the enigmatic hederellids, microconchida, microconchids, and corals. Lily-like crinoids (animals, their resemblance to flowers notwithstanding) were abundant, and trilobites were still fairly common. Bivalvia, Bivalves became commonplace in deep water and outer shelf environments. The first ammonites also appeared during or slightly before the early Devonian Period around 400 Ma. Bactritida, 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 (ostracoderms) declined in diversity, while the jawed fish (gnathostomes) simultaneously increased in both the sea and fresh water. Armored placoderms were numerous during the early ages of the Devonian Period and became extinct in the Late Devonian, perhaps because of competition for food against the other fish species. Early cartilaginous (Chondrichthyes) and bony fishes (Osteichthyes) also become diverse and played a large role within the Devonian seas. The first abundant genus of cartilaginous fish, ''Cladoselache'', appeared in the oceans during the Devonian Period. The great diversity of fish around at the time has led to the Devonian being given the name "The Age of Fishes" 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 (Western Australia), Kimberley Basin of northwest
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
, once extended , fringing a Devonian continent. Reefs are generally built by various carbonate-secreting organisms that can erect wave-resistant structures near sea level. Although modern reefs are constructed mainly by corals and calcareous algae, Devonian reefs were either microbial reefs built up mostly by autotrophic cyanobacteria or coral-stromatoporoid reefs built up by coral-like Stromatoporoidea, stromatoporoids and tabulate and Rugosa, 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, Trigonotarbida, trigonotarbids and Myriapoda, myriapods (although arthropods appeared on land much earlier than in the Early Devonian and the existence of fossils such as ''Protichnites'' suggest that amphibious arthropods may have appeared as early as the Cambrian). By far the largest land organism at the beginning of this period was the enigmatic ''Prototaxites'', which was possibly the fruiting body of an enormous fungus, rolled liverwort mat, or another organism of uncertain affinities that stood more than tall, and towered over the low, carpet-like vegetation during the early part of the Devonian. Also, the first possible fossils of insects appeared around 416  Ma, in the Early Devonian. Evidence for the earliest
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
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'') has been advanced.


The greening of land

Many List of Early Devonian land plants, Early Devonian plants did not have true roots or leaves like extant plants, although vascular tissue is observed in many of those plants. Some of the early land plants such as ''Drepanophycus'' likely spread by vegetative growth and spores. The earliest land plants such as ''Cooksonia'' consisted of leafless, Dichotomous branching, dichotomous axes with terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall. Fossils of ''Armoricaphyton, Armoricaphyton chateaupannense'', about 400 million years old, represent the oldest known plants with woody tissue. By the Middle Devonian, shrub-like forests of primitive plants existed: Lycopodiophyta, lycophytes, Equisetales, horsetails, ferns, and progymnosperms evolved. Most of these plants had true roots and leaves, and many were quite tall. The earliest-known trees appeared in the Middle Devonian. These included a lineage of lycopods and another arborescent, woody vascular plant, the Cladoxylopsida, cladoxylopsids and progymnosperm ''Archaeopteris''. These tracheophytes were able to grow to large size on dry land because they had evolved the ability to biosynthesize lignin, which gave them physical rigidity and improved the effectiveness of their vascular system while giving them resistance to pathogens and herbivores. In
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 ...
age, cladoxylopsid trees formed the first forests in Earth history. 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, and Atmosphere of Earth, atmospheric concentrations of carbon dioxide may have dropped. This may have cooled the climate and led to a massive extinction event. (''See''
Late Devonian extinction The Late Devonian mass extinction, also known as the Kellwasser event, was a mass extinction event which occurred around 372 million years ago, at the boundary between the Frasnian and Famennian ages of the Late Devonian period.Racki, 2005McGh ...
).


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

File:Dunkleosteus terrelli 2024 reconstruction.jpg, ''Dunkleosteus'', one of the largest armoured fish ever to roam the planet, lived during the Late Devonian File:Titanichthys clarki (2024).png, ''Titanichthys'', a Planktivore, planktivorous Arthrodira, arthrodire from the
Famennian The Famennian is the later of two faunal stages in the Late Devonian epoch. The most recent estimate for its duration is that it lasted from around 371.1 to 359.3 million years ago. An earlier 2012 estimate, still used by the International Commis ...
of the Cleveland Shale of Ohio. File:Eastmanosteus pustulosus.jpg, Lower jaw of ''Eastmanosteus pustulosus'' from the Middle Devonian of Wisconsin File:Onychodus.jpg, Tooth of the lobe-finned fish ''Onychodus'' from the Middle Devonian of Wisconsin File:Bothriolepis canadensis (2024).png, ''Bothriolepis'', a diverse Antiarchi, antiarch genus that lived from the Mid to Late Devonian. File:Devonianfishes ntm 1905 smit 1929.gif, Shark-like ''Cladoselache'', several Sarcopterygii, lobe-finned fishes, including ''Eusthenopteron'' that was an early marine tetrapodomorph, and the placoderm ''Bothriolepis'' in a painting from 1905 File:Melocrinites nodosus spinosus.jpg, ''Melocrinites nodosus spinosus'', a spiny, stalked crinoid from the Middle Devonian of Wisconsin File:PhacopidDevonian.jpg, Enrolled phacopid trilobite from the Devonian of Ohio File:AuloporaDevonianSilicaShale.jpg, The common tabulate coral ''Aulopora'' from the Middle Devonian of Ohio – view of colony encrusting a
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of 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 e ...
valve File:Tropidoleptus carinatus.jpg, ''Tropidoleptus carinatus'', an orthid brachiopod from the Middle Devonian of New York File:Pleurodictyum americanum Kashong.jpg, ''Pleurodictyum, Pleurodictyum americanum'', a unique Tabulata, Tabulate coral, Kashong Shale, Middle Devonian of New York File:HederelloidSEM.jpg, SEM image of a Hederellid, hederelloid from the Devonian of Michigan (largest tube diameter is 0.75 mm) File:HederellaOH3.jpg, Devonian spiriferid brachiopod from Ohio which served as a host substrate for a colony of hederelloids File:Prototaxites Dawson1888.PNG, ''Prototaxites'', an 8-meter genus of fungus.


Late Devonian extinction

The
Late Devonian extinction The Late Devonian mass extinction, also known as the Kellwasser event, was a mass extinction event which occurred around 372 million years ago, at the boundary between the Frasnian and Famennian ages of the Late Devonian period.Racki, 2005McGh ...
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 "Extinction event#The "Big Five" mass extinctions, Big Five" mass extinctions in Earth's history. The Devonian extinction crisis primarily affected the marine community, and selectively affected shallow warm-water organisms rather than cool-water organisms. The most important group to be affected by this extinction event were the reef-builders of the great Devonian reef systems. Amongst the severely affected marine groups were the brachiopods, trilobites, ammonites, and acritarchs, and the world saw the disappearance of an estimated 96% of vertebrates like
conodont Conodonts, are an extinct group of marine jawless vertebrates belonging to the class Conodonta (from Ancient Greek κῶνος (''kōnos''), meaning " cone", and ὀδούς (''odoús''), meaning "tooth"). They are primarily known from their hard ...
s and Osteichthyes, bony fishes, and all of the ostracoderms and placoderms. Land plants as well as freshwater species, such as our tetrapod ancestors, were relatively unaffected by the Late Devonian extinction event (there is a counterargument that the Devonian extinctions nearly wiped out the tetrapods). The reasons for the Late Devonian extinctions are still unknown, and all explanations remain speculative. Canadian paleontologist Digby McLaren suggested in 1969 that the Devonian extinction events were caused by an asteroid impact. However, while there were Late Devonian collision events (see the Alamo bolide impact), little evidence supports the existence of a large enough Devonian crater.


See also

* . 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 Devonian, Geological periods Geology of Devon, .