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The Carboniferous ( ) 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 geochronol ...
and
system A system is a group of Interaction, interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its environment (systems), environment, is described by its boundaries, ...
of the
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
that spans 60 million years from the end of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
Period million years ago (
Mya Mya may refer to: Brands and product names * Mya (program), an intelligent personal assistant created by Motorola * Mya (TV channel), an Italian Television channel * Midwest Young Artists, a comprehensive youth music program Codes * Burmese ...
), to the beginning of the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
Period, million years ago. The name ''Carboniferous'' means "coal-bearing", from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
'' carbō'' ("
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when dea ...
") and '' ferō'' ("bear, carry"), and refers to the many coal beds formed globally during that time. The first of the modern 'system' names, it was coined by geologists William Conybeare and
William Phillips William Phillips may refer to: Entertainment * William Phillips (editor) (1907–2002), American editor and co-founder of ''Partisan Review'' * William T. Phillips (1863–1937), American author * William Phillips (director), Canadian film-maker ...
in 1822, based on a study of the British rock succession. The Carboniferous is often treated in North America as two geological periods, the earlier Mississippian and the later Pennsylvanian. Terrestrial animal life was well established by the Carboniferous Period.
Tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s (four limbed vertebrates), which had originated from lobe-finned fish during the preceding
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
, became pentadactylous in and diversified during the Carboniferous, including early
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
lineages such as temnospondyls, with the first appearance of
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s, including synapsids (the group to which modern mammals belong) and
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s during the late Carboniferous. The period is sometimes called the Age of Amphibians, during which amphibians became dominant land vertebrates and diversified into many forms including lizard-like, snake-like, and crocodile-like. Insects would undergo a major radiation during the late Carboniferous. Vast swaths of forest covered the land, which would eventually be laid down and become the coal beds characteristic of the Carboniferous
stratigraphy Stratigraphy is a branch of geology concerned with the study of rock (geology), rock layers (Stratum, strata) and layering (stratification). It is primarily used in the study of sedimentary rock, sedimentary and layered volcanic rocks. Stratigrap ...
evident today. The later half of the period experienced
glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate betw ...
s, low sea level, and
mountain building Mountain formation refers to the geological processes that underlie the formation of mountains. These processes are associated with large-scale movements of the Earth's crust (tectonic plates). Folding, faulting, volcanic activity, igneous intr ...
as the continents collided to form Pangaea. A minor marine and terrestrial extinction event, the Carboniferous rainforest collapse, occurred at the end of the period, caused by climate change.


Etymology and history

The term "Carboniferous" had first been used as an adjective by Irish geologist
Richard Kirwan Richard Kirwan, LL.D, FRS, FRSE MRIA (1 August 1733 – 22 June 1812) was an Irish geologist and chemist. He was one of the last supporters of the theory of phlogiston. Kirwan was active in the fields of chemistry, meteorology, and geology ...
in 1799, and later used in a heading entitled "Coal-measures or Carboniferous Strata" by
John Farey Sr. John Farey Sr. (24 September 1766 – 6 January 1826) was an English geologist and writer best known for Farey sequence, a mathematical construct that is named after him. Biography Youth and early career Farey was born on 24 September 1766 at ...
in 1811, becoming an informal term referring to coal-bearing sequences in Britain and elsewhere in Western Europe. Four units were originally ascribed to the Carboniferous, in ascending order, the Old Red Sandstone, Carboniferous Limestone, Millstone Grit and the
Coal Measures In lithostratigraphy, the coal measures are the coal-bearing part of the Upper Carboniferous System. In the United Kingdom, the Coal Measures Group consists of the Upper Coal Measures Formation, the Middle Coal Measures Formation and the Lower Coal ...
. These four units were placed into a formalised Carboniferous unit by William Conybeare and
William Phillips William Phillips may refer to: Entertainment * William Phillips (editor) (1907–2002), American editor and co-founder of ''Partisan Review'' * William T. Phillips (1863–1937), American author * William Phillips (director), Canadian film-maker ...
in 1822, and later into the Carboniferous System by Phillips in 1835. The Old Red Sandstone was later considered Devonian in age. Subsequently, separate stratigraphic schemes were developed in Western Europe, North America, and Russia. The first attempt to build an international timescale for the Carboniferous was during the Eighth International Congress on Carboniferous Stratigraphy and Geology in Moscow in 1975, when all of the modern ICS stages were proposed.


Stratigraphy

The Carboniferous is divided into two subsystems, the lower Mississippian and upper Pennsylvanian, which are sometimes treated as separate geological periods in North American stratigraphy. Stages can be defined globally or regionally. For global stratigraphic correlation, the
International Commission on Stratigraphy The International Commission on Stratigraphy (ICS), sometimes referred to unofficially as the "International Stratigraphic Commission", is a daughter or major subcommittee grade scientific daughter organization that concerns itself with stratigra ...
(ICS) ratify global stages based on a
Global Boundary Stratotype Section and Point A Global Boundary Stratotype Section and Point (GSSP) is an internationally agreed upon reference point on a stratigraphic section which defines the lower boundary of a stage on the geologic time scale. The effort to define GSSPs is conducted b ...
(GSSP) from a single
formation Formation may refer to: Linguistics * Back-formation, the process of creating a new lexeme by removing or affixes * Word formation, the creation of a new word by adding affixes Mathematics and science * Cave formation or speleothem, a secondar ...
(a stratotype) identifying the lower boundary of the stage. The ICS subdivisions from youngest to oldest are as follows:Cohen, K.M., Finney, S.C., Gibbard, P.L. & Fan, J.-X. (2013; updated
The ICS International Chronostratigraphic Chart
Episodes 36: 199-204.


ICS units

The Mississippian was first proposed by
Alexander Winchell Alexander Winchell (December 31, 1824, in North East, New York – February 19, 1891, in Ann Arbor, Michigan) was a United States geologist who contributed to this field mainly as an educator and a popular lecturer and author. His views on evolu ...
, and the Pennsylvanian was proposed by
J. J. Stevenson John James Stevenson FRSE FSA FRIBA (24 August 1831 – 5 May 1908), usually referred to as J. J. Stevenson, was a British architect of the late-Victorian era. Born in Glasgow, he worked in Glasgow, Edinburgh and London. He is particularly asso ...
in 1888, and both were proposed as distinct and independent systems by H. S. Williams in 1881. The Tournaisian was named after the Belgian city of
Tournai Tournai or Tournay ( ; ; nl, Doornik ; pcd, Tornai; wa, Tornè ; la, Tornacum) is a city and municipality of Wallonia located in the province of Hainaut, Belgium. It lies southwest of Brussels on the river Scheldt. Tournai is part of Euromet ...
. It was introduced in scientific literature by Belgian geologist André Hubert Dumont in 1832. The GSSP for the base of the Tournaisian is located at the
La Serre La Serre (; oc, La Sèrra) is a commune in the Aveyron department in southern France. Population The GSSP Golden Spike for the Tournaisian is in La Serre, with the first appearance of the conodont '' Siphonodella sulcata''. In 2006 it wa ...
section in
Montagne Noire The Montagne Noire ( oc, Montanha Negra, known as the 'Black Mountain' in English) is a mountain range in central southern France. It is located at the southwestern end of the Massif Central at the juncture of the Tarn, Hérault and Aude departm ...
, southern France. It is defined by the
first appearance datum First appearance datum (FAD) is a term used by geologists and paleontologists to designate the first appearance of a species in the geologic record. FADs are determined by identifying the geologically oldest fossil discovered, to date, of a particu ...
of the
conodont Conodonts (Greek ''kōnos'', "cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, which ...
''
Siphonodella sulcata ''Siphonodella'' is an extinct genus of conodonts. ''Siphonodella banraiensis'' is from the Late Devonian of Thailand. ''Siphonodella nandongensis'' is from the Early Carboniferous of the Baping Formation in China.New material of the Early Car ...
'', which was ratified in 1990. However, the GSSP was later shown to have issues, with ''Siphonodella sulcata'' being shown to occur 0.45 m below the proposed boundary. The Viséan Stage was introduced by André Dumont in 1832. Dumont named this stage after the city of
Visé Visé (; nl, Wezet, ; wa, Vizé) is a city and municipality of Wallonia, located on the river Meuse in the province of Liège, Belgium. The municipality consists of the following districts: Argenteau, Cheratte, Lanaye, Lixhe, Richelle, ...
in Belgium's
Liège Province Liège (; wa, Lîdje ; nl, Luik ; german: Lüttich ) is the easternmost province of the Wallonia region of Belgium. Liège Province is the only Belgian province that has borders with three countries. It borders (clockwise from the north) the Du ...
. The GSSP for the Visean is located in Bed 83 at the Pengchong section,
Guangxi Guangxi (; ; Chinese postal romanization, alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Guangxi Zhuang Autonomous Region (GZAR), is an Autonomous regions of China, autonomous region of the People's Republic ...
, southern China, which was ratified in 2012. The GSSP for the base of the Viséan is the first appearance datum of fusulinid (an extinct group of forams) '' Eoparastaffella simplex.'' The Serpukhovian Stage was proposed in 1890 by Russian stratigrapher Sergei Nikitin. It is named after the city of
Serpukhov Serpukhov ( rus, Серпухов, p=ˈsʲɛrpʊxəf) is a city in Moscow Oblast, Russia, located at the confluence of the Oka and the Nara Rivers, south from Moscow ( from Moscow Ring Road) on the Moscow—Simferopol highway. The Moscow— T ...
, near
Moscow Moscow ( , US chiefly ; rus, links=no, Москва, r=Moskva, p=mɐskˈva, a=Москва.ogg) is the capital and largest city of Russia. The city stands on the Moskva River in Central Russia, with a population estimated at 13.0 million ...
. The Serpukhovian Stage currently lacks a defined GSSP. The proposed definition for the base of the Serpukhovian is the first appearance of conodont ''
Lochriea ziegleri ''Lochriea'' is an extinct genus of conodonts. Use in stratigraphy The Visean, the second age of the Mississippian, contains four conodont biozones, two of which are named after ''Lochriea'' species: * the zone of ''Lochriea nodosa'' * the ...
.'' The Bashkirian was named after Bashkiria, the then Russian name of the republic of
Bashkortostan The Republic of Bashkortostan or Bashkortostan ( ba, Башҡортостан Республикаһы, Bashqortostan Respublikahy; russian: Республика Башкортостан, Respublika Bashkortostan),; russian: Респу́блик ...
in the southern
Ural Mountains The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through western ...
of Russia. The stage was introduced by Russian stratigrapher Sofia Semikhatova in 1934. The GSSP for the base of the Bashkirian is located at Arrow Canyon in Nevada, USA, which was ratified in 1996. The GSSP for the base of the Bashkirian is defined by the first appearance of the conodont ''Declinognathodus noduliferus''.'''' The Moscovian is named after Moscow, Russia, and was first introduced by Sergei Nikitin in 1890. The Moscovian currently lacks a defined GSSP. The Kasimovian is named after the Russian city of Kasimov, and originally included as part of Nikitin's original 1890 definition of the Moscovian. It was first recognised as a distinct unit by A.P. Ivanov in 1926, who named it the "'' Tiguliferina''" Horizon after a kind of brachiopod. The Kasimovian currently lacks a defined GSSP. The Gzhelian is named after the Russian village of
Gzhel Gzhel is a Russian style of blue and white ceramics which takes its name from the village of Gzhel and surrounding area, where it has been produced since 1802. Overview About thirty villages located southeast of Moscow produce pottery and shi ...
(russian: Гжель), nearby
Ramenskoye Ramensky (masculine), Ramenskaya (feminine), or Ramenskoye (neuter) may refer to: ;People *Johnny Ramensky (1905–1972), Scottish criminal and war hero *Leonty Ramensky (1884–1953), Russian/Soviet ecologist ;Places *Ramensky District Ramensky ...
, not far from Moscow. The name and type locality were defined by Sergei Nikitin in 1890. The base of the Gzhelian currently lacks a defined GSSP. The GSSP for the base of the Permian is located in the Aidaralash River valley near Aqtöbe, Kazakhstan, which was ratified in 1996. The beginning of the stage is defined by the first appearance of the conodont '' Streptognathodus postfusus.''


Regional stratigraphy


North America

In North American stratigraphy, the Mississippian is divided, in ascending order, into the Kinderhookian, Osagean, Meramecian and Chesterian series, while the Pennsylvanian is divided into the Morrowan, Atokan, Desmoinesian, Missourian and Virgilian series. The Kinderhookian is named after the village of Kinderhook, Pike County, Illinois. It corresponds to the lower part of the Tournasian. The Osagean is named after the Osage River in St. Clair County, Missouri. It corresponds to the upper part of the Tournaisian and the lower part of the Viséan. The Meramecian is named after the Meramec Highlands Quarry, located the near the
Meramec River The Meramec River (), sometimes spelled Maramec River, is one of the longest free-flowing waterways in the U.S. state of Missouri, draining Blanc, Caldwell, and Hawk. "Location" while wandering Blanc, Caldwell, and Hawk. "Executive Summary" fr ...
, southwest of St. Louis, Missouri. It corresponds to the mid Viséan. The Chesterian is named after the Chester Group, a sequence of rocks named after the town of Chester, Illinois. It corresponds to the upper Viséan and all of the Serpukhovian. The Morrowan is named after the Morrow Formation located in NW Arkansas, it corresponds to the lower Bashkirian. The Atokan was originally a formation named after the town of Atoka in southwestern Oklahoma. It corresponds to the upper Bashkirian and lower Moscovian The Desmoinesian is named after the
Des Moines Formation The Des Moines Formation is a geologic formation in Illinois Illinois ( ) is a U.S. state, state in the Midwestern United States, Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metr ...
found near the Des Moines River in central Iowa. It corresponds to the middle and upper Moscovian and lower Kasimovian. The Missourian was named at the same time as the Desmoinesian. It corresponds to the middle and upper Kasimovian. The Virgilian is named after the town of
Virgil, Kansas Virgil is a city in Greenwood County, Kansas, Greenwood County, Kansas, United States. As of the 2020 United States census, 2020 census, the population of the city was 48. It is located approximately 8 miles east of the city of Hamilton, Kansas ...
, it corresponds to the Gzhelian.


Europe

The European Carboniferous is divided into the lower
Dinantian Dinantian is the name of a series (stratigraphy), series or Epoch (reference date), epoch from the Lower Carboniferous system (stratigraphy), system in Europe. It can stand for a series of rocks in Europe or the time span in which they were deposit ...
and upper Silesian, the former being named for the Belgian city of
Dinant Dinant () is a City status in Belgium, city and Municipalities in Belgium, municipality of Wallonia located in the Namur Province, province of Namur, Belgium. On the shores of river Meuse, in the Ardennes, it lies south-east of Brussels, south ...
, and the latter for the
Silesia Silesia (, also , ) is a historical region of Central Europe that lies mostly within Poland, with small parts in the Czech Republic and Germany. Its area is approximately , and the population is estimated at around 8,000,000. Silesia is split ...
region of Central Europe. The boundary between the two subdivisions is older than the Mississippian-Pennsylvanian boundary, lying within the lower Serpukhovian. The boundary has traditionally been marked by the first appearance of the ammonoid '' Cravenoceras leion.'' In Europe, the Dinantian is primarily marine, the so-called "Carboniferous Limestone", while the Silesian primarily known for its coal measures. The Dinantian is divided up into two stages, the Tournaisian and Viséan. The Tournaisian is the same length as the ICS stage, but the Viséan is longer, extending into the lower Serpukhovian. The Silesian is divided into three stages, in ascending order, the Namurian, Westphalian, Stephanian. The Autunian, which corresponds to the middle and upper Gzhelian, is considered a part of the overlying Rotliegend. The Namurian is named after the city of
Namur Namur (; ; nl, Namen ; wa, Nameur) is a city and municipality in Wallonia, Belgium. It is both the capital of the province of Namur and of Wallonia, hosting the Parliament of Wallonia, the Government of Wallonia and its administration. Namu ...
in Belgium. It corresponds to the middle and upper Serpukhovian and the lower Bashkirian. The Westphalian is named after the region of
Westphalia Westphalia (; german: Westfalen ; nds, Westfalen ) is a region of northwestern Germany and one of the three historic parts of the state of North Rhine-Westphalia. It has an area of and 7.9 million inhabitants. The territory of the regio ...
in Germany it corresponds to the upper Bashkirian and all but the uppermost Moscovian. The Stephanian is named after the city of
Saint-Étienne Saint-Étienne (; frp, Sant-Etiève; oc, Sant Estève, ) is a city and the prefecture of the Loire department in eastern-central France, in the Massif Central, southwest of Lyon in the Auvergne-Rhône-Alpes region. Saint-Étienne is the t ...
in eastern France. It corresponds to the uppermost Moscovian, the Kasimovian, and the lower Gzhelian.


Palaeogeography

A global drop in
sea level Mean sea level (MSL, often shortened to sea level) is an average surface level of one or more among Earth's coastal bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical datuma standardised g ...
at the end of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
reversed early in the Carboniferous; this created the widespread inland seas and the
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate g ...
deposition of the Mississippian. There was also a drop in south polar temperatures; southern
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
land was glaciated for much of the period, though it is uncertain if the ice sheets were a holdover from the Devonian or not. These conditions apparently had little effect in the deep tropics, where lush swamps, later to become coal, flourished to within 30 degrees of the northernmost
glacier A glacier (; ) is a persistent body of dense ice that is constantly moving under its own weight. A glacier forms where the accumulation of snow exceeds its Ablation#Glaciology, ablation over many years, often Century, centuries. It acquires dis ...
s. Mid-Carboniferous, a drop in sea level precipitated a major marine extinction, one that hit crinoids and
ammonites Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...
especially hard. This sea level drop and the associated
unconformity An unconformity is a buried erosional or non-depositional surface separating two rock masses or strata of different ages, indicating that sediment deposition was not continuous. In general, the older layer was exposed to erosion for an interval o ...
in North America separate the Mississippian Subperiod from the Pennsylvanian Subperiod. This happened about 323 million years ago, at the onset of the
Permo-Carboniferous Glaciation The Permo-Carboniferous refers to the time period including the latter parts of the Carboniferous and early part of the Permian period. Permo-Carboniferous rocks are in places not differentiated because of the presence of transitional fossils, an ...
. The Carboniferous was a time of active mountain-building as the supercontinent Pangaea came together. The southern
continent A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions are commonly regarded as continents. Ordered from largest in area to smallest, these seven ...
s remained tied together in the supercontinent Gondwana, which collided with North America–Europe ( Laurussia) along the present line of eastern North America. This continental collision resulted in the Hercynian orogeny in Europe, and the Alleghenian orogeny in North America; it also extended the newly uplifted Appalachians southwestward as the Ouachita Mountains. In the same time frame, much of present eastern
Eurasian plate The Eurasian Plate is a tectonic plate that includes most of the continent of Eurasia (a landmass consisting of the traditional continents of Europe and Asia), with the notable exceptions of the Indian subcontinent, the Arabian subcontinent and ...
welded itself to Europe along the line of the
Ural Mountains The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through western ...
. Most of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
supercontinent of Pangea was now assembled, although North China (which would collide in the Latest Carboniferous), and
South China South China () is a geographical and cultural region that covers the southernmost part of China. Its precise meaning varies with context. A notable feature of South China in comparison to the rest of China is that most of its citizens are not n ...
continents were still separated from
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
. The Late Carboniferous Pangaea was shaped like an "O". There were two major oceans in the Carboniferous: Panthalassa and
Paleo-Tethys 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 ...
, which was inside the "O" in the Carboniferous Pangaea. Other minor oceans were shrinking and eventually closed: the Rheic Ocean (closed by the assembly of
South South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
and North America), the small, shallow Ural Ocean (which was closed by the collision of Baltica and Siberia continents, creating the
Ural Mountains The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through western ...
), and the
Proto-Tethys Ocean The Proto-Tethys or Theic Ocean was an ancient ocean that existed from the latest Ediacaran to the Carboniferous (550–330 Ma). History of concept The name "Proto-Tethys" has been used inconsistently for several concepts for a supposed predecess ...
(closed by
North China North China, or Huabei () is a List of regions of China, geographical region of China, consisting of the provinces of Beijing, Tianjin, Hebei, Shanxi and Inner Mongolia. Part of the larger region of Northern China (''Beifang''), it lies north ...
collision with
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
/
Kazakhstania Kazakhstania ( kk, Qazaqstaniya), the Kazakh terranes, or the Kazakhstan Block, is a geological region in Central Asia which consists of the area roughly centered on Lake Balkhash, north and east of the Aral Sea, south of the Siberian craton and w ...
).


Climate

Average global temperatures in the Early Carboniferous Period were high: approximately 20 °C (68 °F). However, cooling during the Middle Carboniferous reduced average global temperatures to about 12 °C (54 °F). Atmospheric
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
levels fell during the Carboniferous Period from roughly 8 times the current level in the beginning, to a level similar to today's at the end. The Carboniferous is considered part of the Late Palaeozoic Ice Age, which began in the latest
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
with the formation of small glaciers in Gondwana. During the Tournaisian the climate warmed, before cooling, there was another warm interval during the Viséan, but cooling began again during the early Serpukhovian. At the beginning of the Pennsylvanian around 323 million years ago, glaciers began to form around the
South Pole The South Pole, also known as the Geographic South Pole, Terrestrial South Pole or 90th Parallel South, is one of the two points where Earth's axis of rotation intersects its surface. It is the southernmost point on Earth and lies antipod ...
, which would grow to cover a vast area of Gondwana. This area extended from the southern reaches of the
Amazon basin The Amazon basin is the part of South America drained by the Amazon River and its tributaries. The Amazon drainage basin covers an area of about , or about 35.5 percent of the South American continent. It is located in the countries of Bolivi ...
and covered large areas of
southern Africa Southern Africa is the southernmost subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number of ...
, as well as most of Australia and Antarctica.
Cyclothems In geology, cyclothems are alternating stratigraphy, stratigraphic sequences of Marine (ocean), marine and non-marine sediments, sometimes interbedded with coal seams. Historically, the term was defined by the European coal geologists who worked ...
, which began around 313 million years ago, and continue into the following Permian indicate that the size of the glaciers were controlled by Milankovitch cycles akin to recent
ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gree ...
s, with glacial periods and
interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene in ...
s. Deep ocean temperatures during this time were cold due to the influx of cold bottom waters generated by seasonal melting of the ice cap. The cooling and drying of the climate led to the Carboniferous Rainforest Collapse (CRC) during the late Carboniferous. Tropical rainforests fragmented and then were eventually devastated by climate change.


Rocks and coal

Carboniferous rocks in Europe and eastern North America largely consist of a repeated sequence of
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
,
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
,
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
and
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when dea ...
beds. In North America, the early Carboniferous is largely marine limestone, which accounts for the division of the Carboniferous into two periods in North American schemes. The Carboniferous coal beds provided much of the fuel for power generation during the
Industrial Revolution The Industrial Revolution was the transition to new manufacturing processes in Great Britain, continental Europe, and the United States, that occurred during the period from around 1760 to about 1820–1840. This transition included going f ...
and are still of great economic importance. The large coal deposits of the Carboniferous may owe their existence primarily to two factors. The first of these is the appearance of
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin th ...
tissue and
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
-bearing trees. The
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of the wood fiber
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 rigidity ...
and the bark-sealing, waxy substance suberin variously opposed decay organisms so effectively that dead materials accumulated long enough to fossilise on a large scale. The second factor was the lower sea levels that occurred during the Carboniferous as compared to the preceding
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
Period. This fostered the development of extensive lowland
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s and
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s in North America and Europe. Based on a genetic analysis of mushroom fungi, it was proposed that large quantities of
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin th ...
were buried during this period because animals and decomposing
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
and fungi had not yet
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
enzymes that could effectively digest the resistant phenolic lignin polymers and waxy suberin polymers. They suggest that fungi that could break those substances down effectively only became dominant towards the end of the period, making subsequent coal formation much rarer. The delayed fungal evolution hypothesis is controversial, however, and has been challenged by other researchers, who conclude that a combination of vast depositional systems present on the continents during the formation of Pangaea and widespread humid, tropical conditions were responsible for the high rate of coal formation. The Carboniferous trees made extensive use of lignin. They had bark to wood ratios of 8 to 1, and even as high as 20 to 1. This compares to modern values less than 1 to 4. This bark, which must have been used as support as well as protection, probably had 38% to 58% lignin. Lignin is insoluble, too large to pass through cell walls, too heterogeneous for specific enzymes, and toxic, so that few organisms other than
Basidiomycetes Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basi ...
fungi can degrade it. To oxidize it requires an atmosphere of greater than 5% oxygen, or compounds such as peroxides. It can linger in soil for thousands of years and its toxic breakdown products inhibit decay of other substances. One possible reason for its high percentages in plants at that time was to provide protection from insects in a world containing very effective insect herbivores (but nothing remotely as effective as modern plant eating insects) and probably many fewer protective toxins produced naturally by plants than exist today. As a result, undegraded carbon built up, resulting in the extensive burial of biologically fixed
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
, leading to an increase in
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
levels in the atmosphere; estimates place the peak oxygen content as high as 35%, as compared to 21% today. This oxygen level may have increased
wildfire A wildfire, forest fire, bushfire, wildland fire or rural fire is an unplanned, uncontrolled and unpredictable fire in an area of Combustibility and flammability, combustible vegetation. Depending on the type of vegetation present, a wildfire ...
activity. It also may have promoted
gigantism Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...
of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s, creatures whose size is today limited by their respiratory systems' ability to transport and distribute oxygen at lower atmospheric concentrations. In eastern North America, marine beds are more common in the older part of the period than the later part and are almost entirely absent by the late Carboniferous. More diverse geology existed elsewhere, of course. Marine life is especially rich in crinoids and other
echinoderms An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea li ...
. Brachiopods were abundant.
Trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
s became quite uncommon. On land, large and diverse
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
populations existed. Land
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
included large amphibians.


Life


Plants

Early Carboniferous Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...
land plants, some of which were preserved in
coal ball A coal ball is a type of concretion, varying in shape from an imperfect sphere to a flat-lying, irregular slab. Coal balls were formed in Carboniferous Period swamps and mires, when peat was prevented from being turned into coal by the high am ...
s, were very similar to those of the preceding Late
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
, but new groups also appeared at this time. The main Early Carboniferous plants were the
Equisetales Equisetales is an order of subclass Equisetidae with only one living family, Equisetaceae, containing the genus ''Equisetum'' (horsetails). Classification In the molecular phylogenetic classification of Smith et al. in 2006, Equisetales, in its ...
(horse-tails),
Sphenophyllales Sphenophyllales is an extinct order of articulate land plants and a sister group to the present-day Equisetales (horsetails). They are fossils dating from the Devonian to the Triassic. They were common during the Late Pennsylvanian to Early ...
(scrambling plants),
Lycopodiales The Lycopodiaceae (class Lycopodiopsida, order Lycopodiales) are an old family of vascular plants, including all of the core clubmosses and firmosses, comprising 16 accepted genera and about 400 known species. This family originated about 380 mil ...
(club mosses), Lepidodendrales (scale trees), Filicales (ferns), Medullosales (informally included in the "
seed ferns A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...
", an assemblage of a number of early
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
groups) and the Cordaitales. These continued to dominate throughout the period, but during
late Carboniferous Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
, several other groups,
Cycadophyta Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
(cycads), the
Callistophytales The Callistophytales was an order of mainly scrambling and lianescent plants found in the wetland "coal swamps" of Euramerica and Cathaysia. They were characterised by having bilaterally-symmetrical, non-cupulate ovules attached to the underside ...
(another group of "seed ferns"), and the
Voltziales Voltziales is an extinct order of conifers. The group contains the ancestral lineages from which all modern conifer groups emerged. The earliest Voltzialean conifers appear in the Late Carboniferous ( Pennsylvanian). Modern conifer lineages emer ...
(related to and sometimes included under the
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
), appeared. The Carboniferous lycophytes of the order Lepidodendrales, which are cousins (but not ancestors) of the tiny club-moss of today, were huge trees with trunks 30 meters high and up to 1.5 meters in diameter. These included '' Lepidodendron'' (with its cone called
Lepidostrobus ''Lepidodendron'' is an extinct genus of primitive vascular plants belonging to the family Lepidodendraceae, part of a group of Lycopodiopsida known as scale trees or arborescent lycophytes, related to quillworts and lycopsids (club mosses). T ...
), '' Anabathra'', '' Lepidophloios'' and '' Sigillaria''. The roots of several of these forms are known as Stigmaria. Unlike present-day trees, their
secondary growth In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of ...
took place in the cortex, which also provided stability, instead of the xylem. The Cladoxylopsids were large trees, that were ancestors of ferns, first arising in the Carboniferous. The fronds of some Carboniferous ferns are almost identical with those of living species. Probably many species were
epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
. Fossil ferns and "seed ferns" include '' Pecopteris'', '' Cyclopteris'', ''
Neuropteris ''Neuropteris'' is an extinct seed fern that existed in the Carboniferous period, known only from fossils. Major species include ''Neuropteris loschi''. See also *Coal forest *''Macroneuropteris ''Macroneuropteris'' is a genus of Carbonifer ...
'', ''
Alethopteris ''Alethopteris'' is a prehistoric plant genus of fossil Pteridospermatophyta (seed ferns) that developed in the Carboniferous period (around ). It is in the family Alethopteridaceae. The genus Alethopteris is among the seed ferns ( Pteridosper ...
'', and ''
Sphenopteris ''Sphenopteris'' is a genus of seed ferns containing the foliage of various extinct plants, ranging from the Devonian to Late Cretaceous. One species, ''S. höninghausi'', was transferred to the genus ''Crossotheca'' in 1911. Biology The frond ...
''; '' Megaphyton'' and '' Caulopteris'' were tree ferns. The Equisetales included the common giant form ''
Calamites ''Calamites'' is a genus of extinct arborescent (tree-like) horsetails to which the modern horsetails (genus ''Equisetum'') are closely related. Unlike their herbaceous modern cousins, these plants were medium-sized trees, growing to heights o ...
'', with a trunk diameter of 30 to and a height of up to . '' Sphenophyllum'' was a slender climbing plant with whorls of leaves, which was probably related both to the calamites and the lycopods. ''
Cordaites ''Cordaites'' is an important genus of extinct gymnosperms which grew on wet ground similar to the Everglades in Florida. Brackish water mussels and crustacea are found frequently between the roots of these trees. The fossils are found in rock s ...
'', a tall plant (6 to over 30 meters) with strap-like leaves, was related to the cycads and conifers; the
catkin A catkin or ament is a slim, cylindrical flower cluster (a spike), with inconspicuous or no petals, usually wind-pollinated (anemophilous) but sometimes insect-pollinated (as in ''Salix''). They contain many, usually unisexual flowers, arranged cl ...
-like reproductive organs, which bore ovules/seeds, is called '' Cardiocarpus''. These plants were thought to live in swamps. True coniferous trees (''
Walchia ''Walchia'' is a fossil conifer, cypress-like genus found in upper Pennsylvanian (Carboniferous) and lower Permian (about 310-290 Mya) rocks of Europe and North America. A forest of in-situ Walchia tree-stumps is located on the Northumberland ...
'', of the order Voltziales) appear later in the Carboniferous, and preferred higher drier ground.


Marine invertebrates

In the oceans the marine invertebrate groups are the
Foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
,
corals Corals are marine invertebrates within the class (biology), class Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important C ...
,
Bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
,
Ostracoda Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typical ...
, brachiopods, ammonoids,
hederelloids Hederellids are extinct colonial animals with calcitic tubular branching exoskeletons. They range from the Silurian to the Permian and were most common in the Devonian period. They are more properly known as "hederelloids" because they were origin ...
,
microconchids The order Microconchida is a group of small, spirally-coiled, encrusting fossil "worm" tubes from the class Tentaculita found from the Upper Ordovician to the Middle Jurassic (Bathonian) around the world. They have lamellar calcitic shells, ...
and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s (especially crinoids). The diversity of brachiopods and fusilinid foraminiferans, surged beginning in the Visean, continuing through the end of the Carboniferous, although cephalopod and nektonic conodont diversity declined. For the first time foraminifera take a prominent part in the marine faunas. The large spindle-shaped genus
Fusulina The Fusulinidae is a family of fusulinacean foraminifera from the upper Carboniferous (Lower Pennsylvanian, Morrowan) to the Upper Permian (Guadalupian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was ...
and its relatives were abundant in what is now Russia, China, Japan, North America; other important genera include ''Valvulina'', ''Endothyra'', ''Archaediscus'', and ''Saccammina'' (the latter common in Britain and Belgium). Some Carboniferous genera are still
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
. The first true priapulids appeared during this period. The microscopic shells of radiolarians are found in
chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a prec ...
s of this age in the Culm of
Devon Devon ( , historically known as Devonshire , ) is a ceremonial and non-metropolitan county in South West England. The most populous settlement in Devon is the city of Plymouth, followed by Devon's county town, the city of Exeter. Devon is ...
and
Cornwall Cornwall (; kw, Kernow ) is a historic county and ceremonial county in South West England. It is recognised as one of the Celtic nations, and is the homeland of the Cornish people. Cornwall is bordered to the north and west by the Atlantic ...
, and in Russia, Germany and elsewhere.
Sponges Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...
are known from spicules and anchor ropes, and include various forms such as the Calcispongea ''Cotyliscus'' and ''Girtycoelia'', the
demosponge Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a har ...
''Chaetetes'', and the genus of unusual colonial glass sponges '' Titusvillia''. Both
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 processes— deposition of sand, wave erosion planing down rock out ...
-building and solitary corals diversify and flourish; these include both
rugose Rugose means "wrinkled". It may refer to: * Rugosa, an extinct order of coral, whose rugose shape earned it the name * Rugose, adjectival form of rugae Species with "rugose" in their names * '' Idiosoma nigrum'', more commonly, a black rugose tr ...
(for example, '' Caninia'', ''Corwenia'', ''Neozaphrentis''), heterocorals, and tabulate (for example, ''Chladochonus'', ''Michelinia'') forms.
Conularids Conulariida is a poorly understood fossil group that has possible affinity with the Cnidaria. Their exact position as a taxon of extinct medusozoan cnidarians is highly speculative. Members of the Conulariida are commonly referred to as conularii ...
were well represented by ''Conularia''
Bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
are abundant in some regions; the fenestellids including ''Fenestella'', ''Polypora'', and ''
Archimedes Archimedes of Syracuse (;; ) was a Greek mathematician, physicist, engineer, astronomer, and inventor from the ancient city of Syracuse in Sicily. Although few details of his life are known, he is regarded as one of the leading scientists ...
'', so named because it is in the shape of an
Archimedean screw The Archimedes screw, also known as the Archimedean screw, hydrodynamic screw, water screw or Egyptian screw, is one of the earliest hydraulic machines. Using Archimedes screws as water pumps (Archimedes screw pump (ASP) or screw pump) dates back ...
. Brachiopods are also abundant; they include productids, some of which reached very large for brachiopods size and had very thick shells (for example, the -wide ''
Gigantoproductus ''Gigantoproductus'' is a genus of extinct brachiopods in the order Productida and the family Monticuliferidae. The species were the largest of the carboniferous brachiopods, with the largest known species reaching in shell width. Such huge ...
''), while others like ''
Chonetes ''Chonetes'' is an extinct genus of brachiopods. It ranged from the Late Ordovician to the Middle Jurassic. Species The following species of ''Chonetes'' have been described: * ''C. (Paeckelmannia)'' * ''C. baragwanathi'' * ''C. billingsi' ...
'' were more conservative in form. Athyridids, spiriferids, rhynchonellids, and terebratulids are also very common. Inarticulate forms include ''
Discina DisCina was a French film production and distribution company established in 1938 by Michel Safra and André Paulvé. It reached its peak during the 1940s and early 1950s, remaining active during the Occupation of France. It also operated a dis ...
'' and '' Crania''. Some species and genera had a very wide distribution with only minor variations.
Annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s such as ''Serpulites'' are common fossils in some horizons. Among the mollusca, the
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
s continue to increase in numbers and importance. Typical genera include ''
Aviculopecten ''Aviculopecten'' is an extinct genus of bivalve mollusc that lived from the Early Devonian to the Late Triassic in Asia, Australia, Europe, North America, and South America.Posidonomya The Posidonia Shale (german: Posidonienschiefer, also called Schistes Bitumineux in Luxembourg) geologically known as the Sachrang Formation, is an Early Jurassic (Toarcian) geological formation of southwestern and northeast Germany, northern Swit ...
'', '' Nucula'', '' Carbonicola'', ''Edmondia'', and ''Modiola''.
Gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s are also numerous, including the genera ''Murchisonia'', ''
Euomphalus ''Euomphalus'' is a genus of fossil marine gastropods known to have lived from the Silurian to the Middle Permian.J.B.Knight,et al 1960. Treatise on Invertebrate Paleontology, Part I, Mollusca 1, ch on systematic descriptions. Geol Soc of Amer ...
'', ''Naticopsis''. Nautiloid
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s are represented by tightly coiled nautilids, with straight-shelled and curved-shelled forms becoming increasingly rare. Goniatite ammonoids such as
Aenigmatoceras ''Aenigmatoceras'' is a genus of ammonite cephalopods from the Carboniferous of Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, N ...
are common.
Trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
s are rarer than in previous periods, on a steady trend towards extinction, represented only by the proetid group.
Ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...
a, a class of
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s, were abundant as representatives of the
meiobenthos Meiobenthos, also called meiofauna, are small benthic invertebrates that live in both marine and fresh water environments. The term ''meiofauna'' loosely defines a group of organisms by their size, larger than microfauna but smaller than macrofau ...
; genera included ''Amphissites'', ''Bairdia'', ''Beyrichiopsis'', ''Cavellina'', ''Coryellina'', ''Cribroconcha'', ''Hollinella'', ''Kirkbya'', ''Knoxiella'', and ''Libumella''. Amongst the
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s, the crinoids were the most numerous. Dense submarine thickets of long-stemmed crinoids appear to have flourished in shallow seas, and their remains were consolidated into thick beds of rock. Prominent genera include ''Cyathocrinus'', ''Woodocrinus'', and ''Actinocrinus''. Echinoids such as ''
Archaeocidaris ''Archaeocidaris'' is an extinct genus of echinoid that lived from the Late Devonian to the Late Permian. Its remains have been found in Africa, Europe, and North America. Sources * ''Fossils'' (Smithsonian Handbooks) by David Ward (Page 1 ...
'' and ''Palaeechinus'' were also present. The
blastoid Blastoids (class Blastoidea) are an extinct type of stemmed echinoderm, often referred to as sea buds. They first appear, along with many other echinoderm classes, in the Ordovician period, and reached their greatest diversity in the Mississi ...
s, which included the Pentreinitidae and Codasteridae and superficially resembled crinoids in the possession of long stalks attached to the seabed, attain their maximum development at this time. File:Aviculopecten subcardiformis01.JPG, ''Aviculopecten subcardiformis''; a
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
from the
Logan Formation The Logan Formation is the name given to a Lower Carboniferous (early Osagean) siltstone, sandstone and conglomeratic unit exposed in east-central Ohio and parts of western West Virginia, USA. Stratigraphy and paleoenvironment The Logan Format ...
(Lower Carboniferous) of
Wooster, Ohio Wooster ( ) is a city in the U.S. state of Ohio and the county seat of Wayne County. Located in northeastern Ohio, the city lies approximately south-southwest of Cleveland, southwest of Akron and west of Canton. The population was 27,232 at t ...
(external mold) File:LoganFauna011312.jpg, Bivalves (''Aviculopecten'') and brachiopods (''Syringothyris'') in the Logan Formation (Lower Carboniferous) in Wooster, Ohio File:Syringothyris01.JPG, ''Syringothyris'' sp.; a spiriferid brachiopod from the Logan Formation (Lower Carboniferous) of Wooster, Ohio (internal mold) File:Palaeophycus01.JPG, ''Palaeophycus'' ichnosp.; a trace fossil from the Logan Formation (Lower Carboniferous) of Wooster, Ohio File:PlatyceratidMississippian.JPG, Crinoid calyx from the Lower Carboniferous of Ohio with a conical platyceratid gastropod (''Palaeocapulus acutirostre'') attached File:Conulariid03.jpg, Conulariid from the Lower Carboniferous of Indiana File:Syringoporid.jpg, Tabulate coral (a syringoporid); Boone Limestone (Lower Carboniferous) near Hiwasse, Arkansas


Freshwater and lagoonal invertebrates

Freshwater Carboniferous invertebrates include various
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s that lived in brackish or fresh water, such as ''Anthraconaia'', ''
Naiadites ''Naiadites'' is an extinct genus of thin-shelled non-marine bivalve from the Upper Carboniferous coal-measures of Nova Scotia in Canada. Species The genus contains the following species: *''Naiadites carbonarius'' from the Pennsylvanian (Upper ...
'', and '' Carbonicola''; diverse
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s such as ''
Candona ''Candona'' is a genus of ostracods in the family Candonidae. Taxonomy The following species are recognised in the genus ''Candona:'' *'' Candona artesensis'' *'' Candona chusanhai'' *'' Candona condensa'' *'' Candona elliptica'' *'' Can ...
'', ''
Carbonita Carbonita is a municipality in the northeast of the Brazilian state of Minas Gerais. the population was 9,414 in a total area of 1,454 km2. The elevation of the town center is 751 meters. It is part of the IBGE statistical meso-region of Jeq ...
'', ''
Darwinula ''Darwinula'' is a genus of ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several ...
'', '' Estheria'', ''Acanthocaris'', ''Dithyrocaris'', and ''Anthrapalaemon''. The eurypterids were also diverse, and are represented by such genera as ''
Adelophthalmus ''Adelophthalmus'' is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of ''Adelophthalmus'' have been discovered in deposits ranging in age from the Early Devonian to the Early Permian, which makes it the longest lived of ...
'', ''
Megarachne ''Megarachne'' is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of ''Megarachne'' have been discovered in deposits of Late Carboniferous age, from the Gzhelian stage, in San Luis, Argentina. The fossils of the single and ...
'' (originally misinterpreted as a giant spider, hence its name) and the specialised very large ''
Hibbertopterus ''Hibbertopterus'' is a genus of eurypterid, a group of extinct aquatic arthropods. Fossils of ''Hibbertopterus'' have been discovered in deposits ranging from the Devonian period in Belgium, Scotland and the United States to the Carboniferous ...
''. Many of these were amphibious. Frequently a temporary return of marine conditions resulted in marine or brackish water genera such as '' Lingula'', ''Orbiculoidea'', and ''
Productus ''Productus subaculeatus'' is an extinct species of brachiopods. Its fossils are present in the Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, ...
'' being found in the thin beds known as marine bands.


Terrestrial invertebrates

Fossil remains of air-breathing
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, myriapods and arachnids are known from the late Carboniferous, but so far not from the early Carboniferous. Their diversity when they do appear, however, shows that these arthropods were both well-developed and numerous. Their large size can be attributed to the moistness of the environment (mostly swampy fern forests) and the fact that the oxygen concentration in the Earth's atmosphere in the Carboniferous was much higher than today. This required less effort for respiration and allowed
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
to grow larger with the up to millipede-like '' Arthropleura'' being the largest-known land invertebrate of all time. Among the insect groups are the huge predatory Protodonata (griffinflies), among which was '' Meganeura'', a giant
dragonfly A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
-like insect and with a wingspan of ca. —the largest flying insect ever to roam the planet. Further groups are the Syntonopterodea (relatives of present-day
mayflies Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are aquatic insects belonging to the orde ...
), the abundant and often large sap-sucking
Palaeodictyopteroidea The Palaeodictyopteroidea or Paleodictyopterida are an extinct superorder of Palaeozoic beaked insects, characterised by unique mouthparts consisting of 5 stylets. They represent the first important terrestrial herbivores, and the first major gr ...
, the diverse herbivorous
Protorthoptera The Protorthoptera are an extinct order of Palaeozoic insects, and represent a wastebasket taxon and paraphyletic assemblage of basal neoptera. They appear during the Middle Carboniferous (late Serpukhovian or early Bashkirian), making them among ...
, and numerous basal
Dictyoptera Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the ord ...
(ancestors of
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
es). Many insects have been obtained from the coalfields of
Saarbrücken Saarbrücken (; french: link=no, Sarrebruck ; Rhine Franconian: ''Saarbrigge'' ; lb, Saarbrécken ; lat, Saravipons, lit=The Bridge(s) across the Saar river) is the capital and largest city of the state of Saarland, Germany. Saarbrücken is S ...
and
Commentry Commentry (; Auvergnat: ''Comentriac'') is a commune in the department of Allier in central France. It lies southwest of Moulins in the valley of the Œil. It is within 8 km of one of the geographic centres of France. The film actress Y ...
, and from the hollow trunks of fossil trees in Nova Scotia. Some British coalfields have yielded good specimens: ''Archaeoptilus'', from the Derbyshire coalfield, had a large wing with preserved part, and some specimens ('' Brodia'') still exhibit traces of brilliant wing colors. In the Nova Scotian tree trunks land snails (''Archaeozonites'', ''Dendropupa'') have been found. File:Meganeura monyi au Museum de Toulouse.jpg, The late Carboniferous giant dragonfly-like insect '' Meganeura'' grew to wingspans of . File:20210116 Pulmonoscorpius kirktonensis.png, The gigantic ''
Pulmonoscorpius ''Pulmonoscorpius'' is an extinct genus of scorpion from the Mississippian (Early Carboniferous) of Scotland. It contains a single named species, ''Pulmonoscorpius kirktonensis''. It was one of the largest scorpions to have ever lived, with th ...
'' from the early Carboniferous reached a length of up to . File:20211224 Arthropleura armata diagrammatic reconstruction.png, ''Arthropleura'' was a giant millipede that fed on the Carboniferous plants. File:Mazothairos1.jpg, ''
Mazothairos ''Mazothairos'' (from Mazo, derived from its location of Mazon Creek and θαιρός, ''thairos'', meaning 'hinge' in Greek) is an extinct genus of very large insect from the Carboniferous period. It was a member of the order Palaeodictyoptera ...
'' was a large
palaeodictyoptera The Palaeodictyoptera are an extinct order of medium-sized to very large, primitive Palaeozoic paleopterous insects. They are informative about the evolution of wings in insects. Overview They were characterized by beak-like mouthparts, used ...
n insect from
Mazon Creek The Mazon Creek fossil beds are a conservation ' found near Morris, in Grundy County, Illinois. The fossils are preserved in ironstone concretions, formed approximately in the mid- Pennsylvanian epoch of the Carboniferous period. These concreti ...
. File:Helenodora inopinata.jpg, ''Helenodora inopinata'', a Stem-group onychophoran known from
Indiana Indiana () is a U.S. state in the Midwestern United States. It is the 38th-largest by area and the 17th-most populous of the 50 States. Its capital and largest city is Indianapolis. Indiana was admitted to the United States as the 19th s ...
File:Blattoidae - Fossil.JPG, A Blattoid
Cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
found in Carboniferous rocks of France


Fish

Many fish inhabited the Carboniferous seas; predominantly Elasmobranchs (sharks and their relatives). These included some, like '' Psammodus'', with crushing pavement-like teeth adapted for grinding the shells of brachiopods, crustaceans, and other marine organisms. Other sharks had piercing teeth, such as the
Symmoriida Symmoriiformes is an extinct order of holocephalians. Originally named Symmoriida by Zangerl (1981), it has subsequently been known by several other names. Lund (1986) synonymized the group with Cladodontida, while Maisey (2008) corrected the na ...
; some, the
petalodont Petalodontiformes ("thin-plate teeth") is an extinct order of marine cartilaginous fish related to modern day chimaera found in what is now the United States of America and Europe.Lund, Richard, E. D. Grogan, and M. Fath. "On the relationships ...
s, had peculiar cycloid cutting teeth. Most of the sharks were marine, but the Xenacanthida invaded fresh waters of the coal swamps. Among the
bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
, the
Palaeonisciformes The Palaeonisciformes (Palaeoniscida) are an extinct order of early ray-finned fishes ( Actinopterygii). Palaeonisciformes ''sensu lato'' first appeared in the fossil record in the Late Silurian and last appeared in the Late Cretaceous. The na ...
found in coastal waters also appear to have migrated to rivers. Sarcopterygian fish were also prominent, and one group, the
Rhizodont Rhizodontida is an extinct group of predatory tetrapodomorphs known from many areas of the world from the Givetian through to the Pennsylvanian - the earliest known species is about 377 million years ago (Mya), the latest around 310 Mya. Rhizodo ...
s, reached very large size. Most species of Carboniferous marine fish have been described largely from teeth, fin spines and dermal ossicles, with smaller freshwater fish preserved whole. Freshwater fish were abundant, and include the genera ''
Ctenodus ''Ctenodus'' (from el, κτείς , 'comb' and el, ὀδούς 'tooth') is an extinct genus of prehistoric lungfish and the longest-surviving genus of Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic tha ...
'', ''Uronemus'', '' Acanthodes'', ''Cheirodus'', and ''
Gyracanthus ''Gyracanthus'' (from el, γύρος , 'curved' and el, ἄκανθα , 'spine') is an extinct genus of acanthodian Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. T ...
''. Chondrichthyes (especially the ''Stethacanthids'') underwent a major evolutionary radiation during the Carboniferous. It is believed that this evolutionary radiation occurred because the decline of the
placoderms Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally 'Plate (animal anatomy), plate-skinned') is a Class (biology), class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devoni ...
at the end of the Devonian Period caused many environmental niches to become unoccupied and allowed new organisms to evolve and fill these niches. As a result of the evolutionary radiation Carboniferous sharks assumed a wide variety of bizarre shapes including ''
Stethacanthus ''Stethacanthus'' is an extinct genus of shark-like holocephaliansCoates, M., Gess, R., Finarelli, J., Criswell, K., Tietjen, K. 2016. A symmoriiform chondrichthyan braincase and the origin of chimaeroid fishes. Nature. doi: 10.1038/nature20806 ...
'' which possessed a flat brush-like dorsal fin with a patch of denticles on its top. ''
Stethacanthus ''Stethacanthus'' is an extinct genus of shark-like holocephaliansCoates, M., Gess, R., Finarelli, J., Criswell, K., Tietjen, K. 2016. A symmoriiform chondrichthyan braincase and the origin of chimaeroid fishes. Nature. doi: 10.1038/nature20806 ...
'' unusual fin may have been used in mating rituals. File:Stethacanthus BW.jpg, ''
Akmonistion ''Akmonistion'' is an extinct genus of holocephalian that lived in the Early Carboniferous. The genus contains a single species, ''A. zangerli'', discovered by Stan Wood in 1982. It is distinguished by an unusual enlarged formation of the dorsal ...
'' of the Holocephali order
Symmoriida Symmoriiformes is an extinct order of holocephalians. Originally named Symmoriida by Zangerl (1981), it has subsequently been known by several other names. Lund (1986) synonymized the group with Cladodontida, while Maisey (2008) corrected the na ...
roamed the oceans of the early Carboniferous. File:Falcatus.jpg, ''
Falcatus ''Falcatus'' is an extinct genus of falcatid chondrichthyan which lived during the early Carboniferous Period in Bear Gulch bay in what is now Montana. Description This fish was quite small, only getting to around 25–30 cm or 10-12 i ...
'' was a Carboniferous holocephalan, with a high degree of sexual dimorphism. File:Dracopristis hoffmanorum.png, ''
Dracopristis ''Dracopristis'' is an extinct genus of ctenacanth (a group of shark-like elasmobranchs) that lived around 307 million years ago, during the Pennsylvanian sub-period of the Carboniferous period. The fish had 12 rows of short, squat teeth, and ...
'' was a Ctenacanthiform elasmobranch from the late Carboniferous of
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
. File:Ornithoprion hertwigi.png,
Ornithoprion ''Ornithoprion'' is an extinct genus of eugeneodont holocephalan closely related to '' Caseodus.'' It lived in the Moscovian stage of the Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 m ...
was a small-sized Eugeneodont holocephalan that had an elongated lower jaw. File:Allenypterus montanus (Restoration) (cropped).jpg, ''
Allenypterus ''Allenypterus'' is a genus of a prehistoric lobe-finned fish which lived during the Bashkirian stage of the Late Carboniferous period, 318 million years ago). Fossils have been discovered in Bear Gulch Limestone, Montana, USA The Unit ...
'' was a Coelacanth fish known from the
Bear Gulch Limestone The Bear Gulch Limestone is a limestone-rich geological lens in central Montana, renowned for the quality of its late Mississippian-aged fossils. It is exposed over a number of outcrops northeast of the Big Snowy Mountains, and is often considere ...
in
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
. File:Echinochimaera meltoni (fossil fish) (Bear Gulch Limestone, Upper Mississippian; Montana, USA) (33784748875).jpg, A fossil of ''
Echinochimaera ''Echinochimaera'' ("prickly chimera") is an extinct genus of Chimaera, chimaeriform fish, known from the Lower Carboniferous Bear Gulch Limestone in Montana, United States. It is one of the earliest Chimaeriformes known. Taxonomy The genus' ...
'', a fish known from the
Bear Gulch Limestone The Bear Gulch Limestone is a limestone-rich geological lens in central Montana, renowned for the quality of its late Mississippian-aged fossils. It is exposed over a number of outcrops northeast of the Big Snowy Mountains, and is often considere ...
in Montana File:†Phanerosteon phonax Carboniferous Bear Gulch.jpg, ''
Phanerosteon ''Phanerosteon'' is an extinct genus of prehistoric bony fish. See also * Prehistoric fish * List of prehistoric bony fish A ''list'' is any set of items in a row. List or lists may also refer to: People * List (surname) Organizations ...
'' was a
Bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...
belonging to the extinct order
Palaeonisciformes The Palaeonisciformes (Palaeoniscida) are an extinct order of early ray-finned fishes ( Actinopterygii). Palaeonisciformes ''sensu lato'' first appeared in the fossil record in the Late Silurian and last appeared in the Late Cretaceous. The na ...
. File:Rhizodus.jpg, ''
Rhizodus ''Rhizodus'' (root tooth) is an extinct genus of basal, finned tetrapodomorphs (the group of sarcopterygians that contains modern tetrapods and their extinct relatives). It belonged to Rhizodontida, one of the earliest-diverging tetrapodomorph c ...
'' was a large freshwater
Rhizodont Rhizodontida is an extinct group of predatory tetrapodomorphs known from many areas of the world from the Givetian through to the Pennsylvanian - the earliest known species is about 377 million years ago (Mya), the latest around 310 Mya. Rhizodo ...
sarcopterygian Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...
from Europe and North America. File:Squatinactis NT small.jpg, ''
Squatinactis ''Squatinactis'' is a genus of extinct Elasmobranchii, elasmobranch Chondrichthyes known from the Carboniferous aged Bear Gulch Limestone, Bear gulch limestone in Montana. This fish was discovered in 1974 by Richard Lund. The Holotype, type speci ...
'', a genus of Elasmobranch fish from Montana


Tetrapods

Carboniferous
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s were diverse and common by the middle of the period, more so than they are today; some were as long as 6 meters, and those fully terrestrial as adults had scaly skin. They included a number of basal tetrapod groups classified in early books under the
Labyrinthodont "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consid ...
ia. These had long bodies, a head covered with bony plates and generally weak or undeveloped limbs. The largest were over 2 meters long. They were accompanied by an assemblage of smaller amphibians included under the Lepospondyli, often only about long. Some Carboniferous amphibians were aquatic and lived in rivers (''
Loxomma ''Loxomma'' (meaning “slanting eyes”) is an extinct genus of Loxommatinae and one of the first Carboniferous tetrapods. They were first described in 1862 and further described in 1870 when two more craniums were found. It is mostly associat ...
'', ''
Eogyrinus ''Pholiderpeton'' (from el, φολῐ́δος , 'horny scale' and el, ἑρπετόν , 'creeping thing') is an extinct genus of embolomere amphibian which lived in the Late Carboniferous period ( Bashkirian) of England. The genus was first ...
'', ''
Proterogyrinus ''Proterogyrinus'' is an extinct genus of early tetrapods from the order Embolomeri. Fossil remains of ''Proterogyrinus'' have been found in Scotland, UK, and West Virginia, United States, and date back to the Serpukhovian (mid-Carboniferous per ...
''); others may have been semi-aquatic (''
Ophiderpeton ''Ophiderpeton'' (from el, ὄφῐς , 'snake' and el, ἑρπετόν 'creeper') is an extinct genus of aistopod tetrapodomorphs from the early Carboniferous to the early Permian. Remains of this genus are widespread and were found in Ohio ...
'', '' Amphibamus'', ''
Hyloplesion ''Hyloplesion'' is an extinct genus of microbrachomorph microsaur. It is the type and only genus within the family Hyloplesiontidae. Fossils have been found from the Czech Republic near the towns of Plzeň, Nýřany, and Třemošná, and da ...
'') or terrestrial (''
Dendrerpeton ''Dendrerpeton'' (from el, δένδρον , 'tree' and el, ἑρπετόν , 'creeping thing') is a genus of an extinct group of temnospondyl amphibians. Its fossils have been found primarily in the Joggins Formation of Eastern Canada and in I ...
'', ''
Tuditanus ''Tuditanus'' is an extinct genus of tuditanid microsaur from the Carboniferous, ~ 306 Ma ago. It was of small size, reaching a length of about 24 cm. See also * Prehistoric amphibian * List of prehistoric amphibians This list of preh ...
'', ''
Anthracosaurus ''Anthracosaurus'' is an extinct genus of embolomere, a possible distant relative of reptiles that lived during the Late Carboniferous (around 310 million years ago) in what is now Scotland and England. It was a large, aquatic eel-like predator a ...
''). The Carboniferous Rainforest Collapse slowed the evolution of amphibians who could not survive as well in the cooler, drier conditions. Amniotes, however, prospered due to specific key adaptations. One of the greatest evolutionary innovations of the Carboniferous was the
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
egg, which allowed the laying of eggs in a dry environment, as well as keratinized scales and claws, allowing for the further exploitation of the land by certain
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s. These included the earliest
sauropsid Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
reptiles ('' Hylonomus''), and the earliest known synapsid ('' Archaeothyris''). Synapsids quickly became huge and diversified in the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
, only for their dominance to stop during the
Mesozoic Era The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising ...
. Sauropsids (reptiles, and also, later, birds) also diversified but remained small until the Mesozoic, during which they would dominate the land, as well as the water and sky. Reptiles underwent a major evolutionary radiation in response to the drier climate that preceded the rainforest collapse. By the end of the Carboniferous Period,
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s had already diversified into a number of groups, including several
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
of synapsid pelycosaurs, protorothyridids,
captorhinids Captorhinidae (also known as cotylosaurs) is an extinct family of tetrapods, traditionally considered primitive reptiles, known from the late Carboniferous to the Late Permian. They had a cosmopolitan distribution across Pangea. Description C ...
, saurians and araeoscelids. File:Pederpes22small.jpg, The
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
-like ''
Pederpes ''Pederpes'' ("Peter's Foot") is an extinct genus of early Carboniferous tetrapod, dating from 348 to 347.6 Ma in the Tournaisian age (lower Mississippian). ''Pederpes'' contains one species, ''P. finneyae'', 1 m long. This most basal Carbonife ...
'', the most primitive tetrapod found in the Mississippian, and known from Scotland. File:Hylonomus BW.jpg, '' Hylonomus'', the earliest
sauropsid Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
, appeared in the Pennsylvanian, and is known from the Joggins Formation in Nova Scotia, and possibly
New Brunswick New Brunswick (french: Nouveau-Brunswick, , locally ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. It is the only province with both English and ...
. File:Petrolacosaurus BW.jpg, '' Petrolacosaurus'', the earliest known diapsid reptile, lived during the late Carboniferous. File:Archaeothyris BW.jpg, '' Archaeothyris'' is the oldest known synapsid, and is found in rocks from
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
. File:Coloraderpeton.jpg, ''
Coloraderpeton ''Coloraderpeton'' is an extinct genus of aïstopod tetrapodomorphs within the family Oestocephalidae. ''Coloraderpeton'' is known from the Carboniferous Sangre de Cristo Formation of Colorado, and was initially known from vertebrae, ribs, and ...
'' was a
snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
-like
aïstopod Aistopoda (Greek for " avingnot-visible feet") is an order of highly specialised snake-like stegocephalians known from the Carboniferous and Early Permian of Europe and North America, ranging from tiny forms only , to nearly in length. They fir ...
tetrapodomorph from the late Carboniferous of
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
. File:Crassygyrinus scoticusDB (flipped).jpg, ''Crassygyrinus'' was a carnivorous stem-tetrapod from the
early Carboniferous Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...
of Scotland. File:Microbrachis BW.jpg, ''
Microbrachis ''Microbrachis'' is an extinct genus of lepospondyl amphibian from the Carboniferous Kladno Formation of the Czech Republic. Description ''Microbrachis'' was an elongated, salamander-like creature, about long, with over 40 vertebrae ins ...
'' was a
lepospondyl Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco (''Diplocaulus minumus''), lepospondyls lived from the Early Carboniferous ( Mississippian) to the Early Per ...
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
known from the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
. File:Amphibamus BW.jpg, '' Amphibamus'' was a dissorophoid
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
from the Late Carboniferous of
Illinois Illinois ( ) is a U.S. state, state in the Midwestern United States, Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolita ...
.


Fungi

As plants and animals were growing in size and abundance in this time (for example, '' Lepidodendron''), land
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
diversified further. Marine fungi still occupied the oceans. All modern classes of fungi were present in the Late Carboniferous ( Pennsylvanian Epoch). During the Carboniferous, animals and bacteria had great difficulty with processing the
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 rigidity ...
and
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
that made up the gigantic trees of the period. Microbes had not evolved that could process them. The trees, after they died, simply piled up on the ground, occasionally becoming part of long-running wildfires after a lightning strike, with others very slowly degrading into
coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when dea ...
. White rot fungus were the first organisms to be able to process these and break them down in any reasonable quantity and timescale. Thus, some have proposed that fungi helped end the Carboniferous Period, stopping accumulation of undegraded plant matter, although this idea remains highly controversial.


Extinction events


Romer's gap

The first 15 million years of the Carboniferous had very limited terrestrial fossils. This gap in the fossil record is called
Romer's gap Romer's gap is an example of an apparent gap in the tetrapod fossil record used in the study of evolutionary biology. Such gaps represent periods from which excavators have not yet found relevant fossils. Romer's gap is named after paleontologist ...
after the American palaentologist Alfred Romer. While it has long been debated whether the gap is a result of fossilisation or relates to an actual event, recent work indicates the gap period saw a drop in atmospheric oxygen levels, indicating some sort of
ecological collapse Ecological collapse refers to a situation where an ecosystem suffers a drastic, possibly permanent, reduction in carrying capacity for all organisms, often resulting in mass extinction. Usually, an ecological collapse is precipitated by a disastro ...
. The gap saw the demise of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
fish-like ichthyostegalian labyrinthodonts, and the rise of the more advanced
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
and reptiliomorphan amphibians that so typify the Carboniferous terrestrial vertebrate fauna.


Carboniferous rainforest collapse

Before the end of the Carboniferous Period, an
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 change in the diversity and abundance of multicellular organisms. I ...
occurred. On land this event is referred to as the Carboniferous Rainforest Collapse (CRC). Vast tropical rainforests collapsed suddenly as the climate changed from hot and humid to cool and arid. This was likely caused by intense
glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate betw ...
and a drop in sea levels. The new climatic conditions were not favorable to the growth of rainforest and the animals within them. Rainforests shrank into isolated islands, surrounded by seasonally dry habitats. Towering
lycopsid Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching s ...
forests with a heterogeneous mixture of vegetation were replaced by much less diverse tree-fern dominated flora. Amphibians, the dominant vertebrates at the time, fared poorly through this event with large losses in biodiversity; reptiles continued to diversify due to key adaptations that let them survive in the drier habitat, specifically the hard-shelled egg and scales, both of which retain water better than their amphibian counterparts.


See also

*
Carboniferous tetrapods Carboniferous tetrapods include amphibians and reptiles that lived during the Carboniferous Period. Though stem-tetrapods originated in the preceding Devonian, it was in the earliest Carboniferous that the first crown tetrapods appeared, with ful ...
* Carboniferous Rainforest Collapse * Important Carboniferous Lagerstätten **
East Kirkton Quarry East Kirkton Quarry is a former limestone quarry in West Lothian, Scotland (East Kirkton Limestone), now better known as a fossil site known for terrestrial fossils from the fossil-poor Romer's gap, a 15 million year period at the beginning of th ...
; c. 350 mya;
Bathgate Bathgate ( sco, Bathket or , gd, Both Chèit) is a town in West Lothian, Scotland, west of Livingston and adjacent to the M8 motorway. Nearby towns are Armadale, Blackburn, Linlithgow, Livingston, West Calder and Whitburn. Situated sout ...
, Scotland **
Hamilton Quarry Hamilton Quarry is a Late Carboniferous lagerstätte near Hamilton, Kansas, United States. It has a diverse assemblage of unusually well-preserved marine, euryhaline, freshwater, flying, and terrestrial fossils (invertebrates, vertebrates, and pla ...
; 320 mya;
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the ...
, US **
Mazon Creek The Mazon Creek fossil beds are a conservation ' found near Morris, in Grundy County, Illinois. The fossils are preserved in ironstone concretions, formed approximately in the mid- Pennsylvanian epoch of the Carboniferous period. These concreti ...
; 300 mya;
Illinois Illinois ( ) is a U.S. state, state in the Midwestern United States, Midwestern United States. Its largest metropolitan areas include the Chicago metropolitan area, and the Metro East section, of Greater St. Louis. Other smaller metropolita ...
, US * List of fossil sites ''(with link directory)''


References


Sources

* * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * *


External links

*
Examples of Carboniferous Fossils60+ images of Carboniferous Foraminifera
{{Authority control Geological periods