Devonian
   HOME

TheInfoList



OR:

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 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 earliest land animals, 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. Tetrapodomorphs, which include the ancestors of all four- limbed 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 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 hindlimbs, 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. The subsequent end-Devonian extinction, 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 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 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 was a lengthy debate between Roderick Murchison,
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 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 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 from to Ma and then by the Emsian, which lasted until the Middle Devonian began at Ma. During this time, the first ammonoids 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 and Clymeniida. 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, which then gave way to the Givetian 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, from to Ma, during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing Famennian subdivision, the beginning and end of which are marked with extinction events. This lasted until the end of the Devonian 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 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.


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.


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 Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
. 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 Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
and
Ellesmere Island Ellesmere Island (; ) is Canada's northernmost and List of Canadian islands by area, third largest island, and the List of islands by area, tenth largest in the world. It comprises an area of , slightly smaller than Great Britain, and the total ...
) established tropical conditions, most of the continent was located within the natural dry zone along the
Tropic of Capricorn The Tropic of Capricorn (or the Southern Tropic) is the circle of latitude that contains the subsolar point at the December (or southern) solstice. It is thus the southernmost latitude where the Sun can be seen directly overhead. It also reach ...
, which (as nowadays) is a result of the convergence of two great air-masses, the
Hadley cell The Hadley cell, also known as the Hadley circulation, is a global-scale tropical atmospheric circulation that features air rising near the equator, flowing poleward near the tropopause at a height of above the Earth's surface, cooling and des ...
and the
Ferrel cell Atmospheric circulation is the large-scale movement of air and together with ocean circulation is the means by which thermal energy is redistributed on the surface of the Earth. The Earth's atmospheric circulation varies from year to year, but ...
. In these near-deserts, the Old Red Sandstone sedimentary beds formed, made red by the oxidised iron (
hematite Hematite (), also spelled as haematite, is a common iron oxide compound with the formula, Fe2O3 and is widely found in rocks and soils. Hematite crystals belong to the rhombohedral lattice system which is designated the alpha polymorph of . ...
) characteristic of drought conditions. The abundance of red sandstone on continental land also lends Laurussia the name "the Old Red Continent". For much of the Devonian, the majority of western Laurussia (North America) was covered by subtropical inland seas which hosted a diverse ecosystem of reefs and marine life. Devonian marine deposits are particularly prevalent in the
midwestern The Midwestern United States (also referred to as the Midwest, the Heartland or the American Midwest) is one of the four census regions defined by the United States Census Bureau. It occupies the northern central part of the United States. It ...
and northeastern United States. Devonian reefs also extended along the southeast edge of Laurussia, a coastline now corresponding to southern
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
,
Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
, and other mid-latitude areas of Europe. In the Early and Middle Devonian, the west coast of Laurussia was a passive margin with broad coastal waters, deep silty embayments, river deltas and estuaries, found today in
Idaho Idaho ( ) is a landlocked U.S. state, state in the Pacific Northwest and Mountain states, Mountain West subregions of the Western United States. It borders Montana and Wyoming to the east, Nevada and Utah to the south, and Washington (state), ...
and
Nevada Nevada ( ; ) is a landlocked state in the Western United States. It borders Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the seventh-most extensive, th ...
. In the Late Devonian, an approaching volcanic
island arc Island arcs are long archipelago, chains of active volcanoes with intense earthquake, seismic activity found along convergent boundary, convergent plate tectonics, tectonic plate boundaries. Most island arcs originate on oceanic crust and have re ...
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 Svalbard ( , ), previously known as Spitsbergen or Spitzbergen, is a Norway, Norwegian archipelago that lies at the convergence of the Arctic Ocean with the Atlantic Ocean. North of continental Europe, mainland Europe, it lies about midway be ...
. Many of the Devonian collisions in Laurussia produce both mountain chains and
foreland basin A foreland basin is a structural basin that develops adjacent and parallel to a mountain belt. Foreland basins form because the immense mass created by crustal thickening associated with the evolution of a mountain belt causes the lithospher ...
s, which are frequently fossiliferous.


Gondwana

Gondwana was by far the largest continent on the planet. It was completely south of the equator, although the northeastern sector (now Australia) did reach tropical latitudes. The southwestern sector (now South America) was located to the far south, with
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
situated near the
South Pole The South Pole, also known as the Geographic South Pole or Terrestrial South Pole, is the point in the Southern Hemisphere where the Earth's rotation, Earth's axis of rotation meets its surface. It is called the True South Pole to distinguish ...
. The northwestern edge of Gondwana was an active margin for much of the Devonian, and saw the accretion of many smaller land masses and island arcs. These include Chilenia, Cuyania, and Chaitenia, which now form much of
Chile Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...
and
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
. These collisions were associated with
volcanic activity Volcanism, vulcanism, volcanicity, or volcanic activity is the phenomenon where solids, liquids, gases, and their mixtures erupt to the surface of a solid-surface astronomical body such as a planet or a moon. It is caused by the presence of a he ...
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 Tibet (; ''Böd''; ), or Greater Tibet, is a region in the western part of East Asia, covering much of the Tibetan Plateau and spanning about . It is the homeland of the Tibetan people. Also resident on the plateau are other ethnic groups s ...
. The eastern margin, though warmer than the west, was equally active. Numerous mountain building events and
granite Granite ( ) is a coarse-grained (phanerite, phaneritic) intrusive rock, intrusive igneous rock composed mostly of quartz, alkali feldspar, and plagioclase. It forms from magma with a high content of silica and alkali metal oxides that slowly coo ...
and
kimberlite Kimberlite is an igneous rock and a rare variant of peridotite. It is most commonly known as the main host matrix for diamonds. It is named after the town of Kimberley, Northern Cape, Kimberley in South Africa, where the discovery of an 83.5-Car ...
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 Tasmania (; palawa kani: ''Lutruwita'') is an island States and territories of Australia, state of Australia. It is located to the south of the Mainland Australia, Australian mainland, and is separated from it by the Bass Strait. The sta ...
, and Antarctica.


Asian terranes

Several island microcontinents (which would later coalesce into modern day Asia) stretched over a low-latitude
archipelago An archipelago ( ), sometimes called an island group or island chain, is a chain, cluster, or collection of islands. An archipelago may be in an ocean, a sea, or a smaller body of water. Example archipelagos include the Aegean Islands (the o ...
to the north of Gondwana. They were separated from the southern continent by an oceanic basin: the Paleo-Tethys. Although the western Paleo-Tethys Ocean had existed since the Cambrian, the eastern part only began to rift apart as late as the Silurian. This process accelerated in the Devonian. The eastern branch of the Paleo-Tethys was fully opened when
South China South China ( 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 ...
and 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 Mainland Southeast Asia (historically known as Indochina and the Indochinese Peninsula) is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to th ...
), together as a unified continent, detached from the northeastern sector of Gondwana. Nevertheless, they remained close enough to Gondwana that their Devonian fossils were more closely related to Australian species than to north Asian species. Other Asian terranes remained attached to Gondwana, including Sibumasu (western Indochina), Tibet, and the rest of the
Cimmerian The Cimmerians were an Ancient Iranian peoples, ancient Eastern Iranian languages, Eastern Iranian peoples, Iranic Eurasian nomads, equestrian nomadic people originating in the Pontic–Caspian steppe, part of whom subsequently migrated into W ...
blocks. While the South China-Annamia continent was the newest addition to the Asian microcontinents, it was not the first.
North China North China () is a list of regions of China, geographical region of the People's Republic of China, consisting of five province-level divisions of China, provincial-level administrative divisions, namely the direct-administered municipalities ...
and the Tarim Block (now northwesternmost China) were located westward and continued to drift northwards, powering over older oceanic crust in the process. Further west was a small ocean (the Turkestan Ocean), followed by the larger microcontinents of
Kazakhstania Kazakhstania (), the Kazakh terranes, or the Kazakhstan Block, is a geological region in Central Asia which consists of the area roughly centered on Lake Balkhash, north and east of the Aral Sea, south of the Siberian craton and west of the Alta ...
,
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 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 plume A mantle plume is a proposed mechanism of convection within the Earth's mantle, hypothesized to explain anomalous volcanism. Because the plume head partially melts on reaching shallow depths, a plume is often invoked as the cause of volcanic ho ...
s impacted the continent with
flood basalt A flood basalt (or plateau basalt) is the result of a giant volcanic eruption or series of eruptions that covers large stretches of land or the ocean floor with basalt lava. Many flood basalts have been attributed to the onset of a hotspot (geolo ...
s during the Late Devonian. The
Altai-Sayan region The Altai-Sayan region is an area of Inner Asia proximate to the Altai Mountains and the Sayan Mountains, near to where Russia, China, Mongolia and Kazakhstan come together. This region is one of the world centers of Temperate climate, temperate pl ...
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 Manchuria is a historical region in northeast Asia encompassing the entirety of present-day northeast China and parts of the modern-day Russian Far East south of the Uda (Khabarovsk Krai), Uda River and the Tukuringra-Dzhagdy Ranges. The exact ...
,
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
and their vicinities). Though certainly close to Siberia in the Devonian, the precise location of Amuria is uncertain due to contradictory paleomagnetic data.


Closure of the Rheic Ocean

The Rheic Ocean, which separated Laurussia from Gondwana, was wide at the start of the Devonian, having formed after the drift of
Avalonia Avalonia was a microcontinent in the Paleozoic era. Crustal fragments of this former microcontinent are terranes in parts of the eastern coast of North America: Atlantic Canada, and parts of the East Coast of the United States, East Coast of the ...
away from Gondwana. It steadily shrunk as the period continued, as the two major continents approached near 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 ...
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 Subduction is a geological process in which the oceanic lithosphere and some continental lithosphere is recycled into the Earth's mantle at the convergent boundaries between tectonic plates. Where one tectonic plate converges with a second p ...
and
metamorphism Metamorphism is the transformation of existing Rock (geology), rock (the protolith) to rock with a different mineral composition or Texture (geology), texture. Metamorphism takes place at temperatures in excess of , and often also at elevated ...
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 The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, compri ...
,
Armorica In ancient times, Armorica or Aremorica (Gaulish: ; ; ) was a region of Gaul between the Seine and the Loire that includes the Brittany Peninsula, and much of historical Normandy. Name The name ''Armorica'' is a Latinized form of the Gauli ...
(France), Palaeo-Adria (the western Mediterranean area),
Bohemia Bohemia ( ; ; ) is the westernmost and largest historical region of the Czech Republic. In a narrow, geographic sense, it roughly encompasses the territories of present-day Czechia that fall within the Elbe River's drainage basin, but historic ...
,
Franconia Franconia ( ; ; ) is a geographical region of Germany, characterised by its culture and East Franconian dialect (). Franconia is made up of the three (governmental districts) of Lower Franconia, Lower, Middle Franconia, Middle and Upper Franco ...
, and Saxothuringia. These continental blocks, collectively known as the Armorican Terrane Assemblage, split away from Gondwana in the Silurian and drifted towards Laurussia through the Devonian. Their collision with Laurussia leads to the beginning of the Variscan Orogeny, a major mountain-building event which would escalate further in the Late Paleozoic. Franconia and Saxothuringia collided with Laurussia near the end of the Early Devonian, pinching out the easternmost Rheic Ocean. The rest of the Armorican terranes followed, and by the end of the Devonian they were fully connected with Laurussia. This sequence of rifting and collision events led to the successive creation and destruction of several small seaways, including the Rheno-Hercynian, Saxo-Thuringian, and Galicia-Moldanubian oceans. Their sediments were eventually compressed and completely buried as Gondwana fully collided with Laurussia in the Carboniferous.


Life


Marine biota

Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by
conodonts Conodonts, are an extinct group of marine jawless vertebrates belonging to the Class (biology), class Conodonta (from Ancient Greek κῶνος (''kōnos''), meaning "cone", and ὀδούς (''odoús''), meaning "tooth"). They are primarily known ...
,
bryozoans Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
, 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, microconchids, and
coral Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
s. Lily-like
crinoid Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in their adult form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are ...
s (animals, their resemblance to flowers notwithstanding) were abundant, and
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 were still fairly common.
Bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
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. 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 Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salt (chemistry), salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include ...
. 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 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 ...
) and bony fishes (
Osteichthyes Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
) also become diverse and played a large role within the Devonian seas. The first abundant genus of cartilaginous fish, ''
Cladoselache ''Cladoselache'' ("branch shark") is an Extinction, extinct genus of shark-like chondrichthyan (cartilaginous fish) from the Late Devonian (Famennian) of North America. It was similar in body shape to modern Lamnidae, lamnid sharks (such as Isuru ...
'', 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 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 A carbonate is a salt of carbonic acid, (), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word "carbonate" may also refer to a carbonate ester, an organic compound containing the carbonate group ...
-secreting organisms that can erect wave-resistant structures near sea level. Although modern reefs are constructed mainly by corals and calcareous
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
, Devonian reefs were either microbial reefs built up mostly by
autotrophic An autotroph is an organism that can convert abiotic sources of energy into energy stored in organic compounds, which can be used by other organisms. Autotrophs produce complex organic compounds (such as carbohydrates, fats, and proteins) us ...
cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
or coral-stromatoporoid reefs built up by coral-like stromatoporoids and tabulate and rugose corals. Microbial reefs dominated under the warmer conditions of the early and late Devonian, while coral-stromatoporoid reefs dominated during the cooler middle Devonian.


Terrestrial biota

By the Devonian Period, life was well underway in its colonization of the land. The
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
forests and
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
l and algal mats of the Silurian were joined early in the period by primitive rooted plants that created the first stable
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
s and harbored arthropods like
mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
s,
scorpion Scorpions are predatory arachnids of the Order (biology), order Scorpiones. They have eight legs and are easily recognized by a pair of Chela (organ), grasping pincers and a narrow, segmented tail, often carried in a characteristic forward cur ...
s, trigonotarbids and myriapods (although arthropods appeared on land much earlier than in the Early Devonian and the existence of fossils such as '' Protichnites'' suggest that amphibious arthropods may have appeared as early as the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
). 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
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s 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 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 A dichotomy () is a partition of a set, partition of a whole (or a set) into two parts (subsets). In other words, this couple of parts must be * jointly exhaustive: everything must belong to one part or the other, and * mutually exclusive: nothi ...
axes with terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall. Fossils of '' Armoricaphyton chateaupannense'', about 400 million years old, represent the oldest known plants with
wood Wood is a structural tissue/material found as xylem in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulosic fibers that are strong in tension and embedded in a matrix of lignin t ...
y tissue. By the Middle Devonian, shrub-like forests of primitive plants existed: lycophytes, horsetails,
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s, 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 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 Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidit ...
, which gave them physical rigidity and improved the effectiveness of their vascular system while giving them resistance to pathogens and herbivores. In Eifelian 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 A carbon sink is a natural or artificial carbon sequestration process that "removes a  greenhouse gas, an aerosol or a precursor of a greenhouse gas from the atmosphere". These sinks form an important part of the natural carbon cycle. An overar ...
, and atmospheric concentrations of
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
may have dropped. This may have cooled the climate and led to a massive
extinction event An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp fall in the diversity and abundance of multicellular organisms. It occ ...
. (''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 Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...
and sediment deposition. The rapid evolution of a terrestrial ecosystem that contained copious animals opened the way for the first
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s 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 planktivorous
arthrodire Arthrodira (Greek for "jointed neck") is an Order (biology), order of extinct armored, jawed fishes of the class Placodermi that flourished in the Devonian period before their sudden extinction, surviving for about 50 million years and penetratin ...
from the Famennian of the Cleveland Shale of
Ohio Ohio ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Erie to the north, Pennsylvania to the east, West Virginia to the southeast, Kentucky to the southwest, Indiana to the ...
. 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 antiarch
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
that lived from the Mid to Late Devonian. File:Devonianfishes ntm 1905 smit 1929.gif, Shark-like ''
Cladoselache ''Cladoselache'' ("branch shark") is an Extinction, extinct genus of shark-like chondrichthyan (cartilaginous fish) from the Late Devonian (Famennian) of North America. It was similar in body shape to modern Lamnidae, lamnid sharks (such as Isuru ...
'', several 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 Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in their adult form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are ...
from the Middle Devonian of
Wisconsin Wisconsin ( ) is a U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest of the United States. It borders Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake Michigan to the east, Michig ...
File:PhacopidDevonian.jpg, Enrolled phacopid
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 ...
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 americanum'', a unique Tabulate coral, Kashong Shale, Middle Devonian of New York File:HederelloidSEM.jpg, SEM image of a hederelloid from the Devonian of Michigan (largest tube diameter is 0.75 mm) File:HederellaOH3.jpg, Devonian spiriferid brachiopod from
Ohio Ohio ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Erie to the north, Pennsylvania to the east, West Virginia to the southeast, Kentucky to the southwest, Indiana to the ...
which served as a host substrate for a colony of hederelloids File:Prototaxites Dawson1888.PNG, '' Prototaxites'', an 8-meter genus of
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
.


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 " 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 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 Geological periods .