The CARBONIFEROUS is a geologic period and system that spans 60
million years from the end of the
Devonian Period 358.9 million years
ago (Mya ), to the beginning of the
Permian Period, 298.9 Mya. The
Carboniferous means "coal-bearing" and derives from the Latin
words carbō ("coal ") and ferō ("I bear, I carry"), and was coined
by geologists William Conybeare and William Phillips in 1822.
Based on a study of the British rock succession, it was the first of
the modern 'system' names to be employed, and reflects the fact that
many coal beds were formed globally during that time. The
Carboniferous is often treated in
North America as two geological
periods, the earlier Mississippian and the later Pennsylvanian .
Terrestrial life was well established by the
Amphibians were the dominant land vertebrates, of which one branch
would eventually evolve into amniotes , the first solely terrestrial
Arthropods were also very common, and many (such as
Meganeura ) were
much larger than those of today. Vast swaths of forest covered the
land, which would eventually be laid down and become the coal beds
characteristic of the
Carboniferous stratigraphy evident today. The
atmospheric content of oxygen also reached their highest levels in
geological history during the period, 35% compared with 21% today,
allowing terrestrial invertebrates to evolve to great size.
A major marine and terrestrial extinction event, the Carboniferous
rainforest collapse , occurred in the middle of the period, caused by
climate change. The later half of the period experienced glaciations
, low sea level, and mountain building as the continents collided to
* 1 Subdivisions
* 2 Palaeogeography
* 3 Climate
* 4 Rocks and coal
* 5 Life
* 5.1 Plants
* 5.2 Marine invertebrates
* 5.3 Freshwater and lagoonal invertebrates
* 5.4 Terrestrial invertebrates
* 5.5 Fish
* 5.6 Tetrapods
* 6 Extinction events
* 6.1 Romer\'s gap
Carboniferous rainforest collapse
* 7 See also
* 8 References
* 9 Sources
* 10 External links
United States the
Carboniferous is usually broken into
Mississippian (earlier) and Pennsylvanian (later) subperiods. The
Mississippian is about twice as long as the Pennsylvanian, but due to
the large thickness of coal-bearing deposits with Pennsylvanian ages
in Europe and North America, the two subperiods were long thought to
have been more or less equal in duration. In Europe the Lower
Carboniferous sub-system is known as the
Dinantian , comprising the
Visean Series, dated at 362.5-332.9 Ma, and the Upper
Carboniferous sub-system is known as the Silesian , comprising the
Namurian , Westphalian , and Stephanian Series, dated at 332.9-298.9
Ma. The Silesian is roughly contemporaneous with the late
Serpukhovian plus the Pennsylvanian. In Britain the
Dinantian is traditionally known as the
Carboniferous Limestone , the
Namurian as the
Millstone Grit , and the Westphalian as the Coal
Pennant Sandstone .
The faunal stages from youngest to oldest, together with some of
their subdivisions, are:
LATE PENNSYLVANIAN: GZHELIAN (MOST RECENT)
* Noginskian / Virgilian (part)
LATE PENNSYLVANIAN: KASIMOVIAN
* Dorogomilovksian / Virgilian (part)
* Chamovnicheskian / Cantabrian / Missourian
* Krevyakinskian / Cantabrian / Missourian
MIDDLE PENNSYLVANIAN: MOSCOVIAN
* Myachkovskian / Bolsovian / Desmoinesian
* Podolskian / Desmoinesian
* Kashirskian / Atokan
* Vereiskian / Bolsovian / Atokan
EARLY PENNSYLVANIAN: BASHKIRIAN / MORROWAN
* Melekesskian / Duckmantian
* Cheremshanskian / Langsettian
LATE MISSISSIPPIAN: SERPUKHOVIAN
* Chokierian / Chesterian / Elvirian
* Arnsbergian / Elvirian
MIDDLE MISSISSIPPIAN: VISEAN
* Brigantian / St Genevieve / Gasperian / Chesterian
* Asbian / Meramecian
* Holkerian / Salem
* Arundian / Warsaw / Meramecian
* Chadian / Keokuk / Osagean (part) / Osage (part)
EARLY MISSISSIPPIAN: TOURNAISIAN (OLDEST)
* Ivorian / (part) / Osage (part)
* Hastarian / Kinderhookian / Chouteau
A global drop in sea level at the end of the
Devonian reversed early
in the Carboniferous; this created the widespread inland seas and the
carbonate deposition of the Mississippian. There was also a drop in
south polar temperatures; southern Gondwanaland was glaciated
throughout 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 glaciers .
Generalized geographic map of the
United States in Middle
Mid-Carboniferous, a drop in sea level precipitated a major marine
extinction, one that hit crinoids and ammonites especially hard. This
sea level drop and the associated unconformity in North America
separate the Mississippian subperiod from the Pennsylvanian subperiod.
This happened about 323 million years ago, at the onset of the
Carboniferous was a time of active mountain-building , as the
Pangaea came together. The southern continents 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 welded itself to Europe along the line of the Ural
mountains . Most of the
Mesozoic supercontinent of Pangea was now
assembled, although North China (which would collide in the Latest
Carboniferous), and South China continents were still separated from
Laurasia . The Late
Pangaea was shaped like an "O."
There were two major oceans in the Carboniferous—
Paleo-Tethys , which was inside the "O" in the
Other minor oceans were shrinking and eventually closed - Rheic Ocean
(closed by the assembly of South and
North America ), the small,
Ural Ocean (which was closed by the collision of
Siberia continents, creating the
Ural Mountains ) and Proto-Tethys
Ocean (closed by North China collision with Siberia /
Average global temperatures in the Early
Carboniferous Period were
high: approximately 20 °C (68 °F). However, cooling during the
Carboniferous reduced average global temperatures to about 12
°C (54 °F). Lack of growth rings of fossilized trees suggest a lack
of seasons of a tropical climate. Glaciations in
Gondwana , triggered
by Gondwana's southward movement, continued into the
because of the lack of clear markers and breaks, the deposits of this
glacial period are often referred to as
Permo-Carboniferous in age.
The thicker atmosphere and stronger coriolis effect due to Earth's
faster rotation (a day lasted for 22.4 hours in early Carboniferous)
created significantly stronger winds than today.
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
ROCKS AND COAL
Carboniferous marble in Big Cottonwood Canyon, Wasatch
Carboniferous rocks in Europe and eastern
North America largely
consist of a repeated sequence of limestone , sandstone , shale and
coal 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
beds provided much of the fuel for power generation during the
Industrial Revolution 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
tissue and bark -bearing trees. The evolution of the wood fiber lignin
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
Devonian period. This promoted the development of extensive
lowland swamps and forests in
North America and Europe. Based on a
genetic analysis of mushroom fungi, it was proposed that large
quantities of wood were buried during this period because animals and
decomposing bacteria had not yet evolved 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.
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
Basidiomycetes 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 insectivores )
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 , leading to an
increase in oxygen 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 activity. It also may have promoted
gigantism of insects and amphibians — creatures that have been
constrained in size by respiratory systems that are limited in their
physiological ability to transport and distribute oxygen at the lower
atmospheric concentrations that have since been available.
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 . Brachiopods were abundant. Trilobites became quite
uncommon. On land, large and diverse plant populations existed. Land
vertebrates included large amphibians.
Etching depicting some of the most significant plants of the
Wikisource has the text of the 1879
American Cyclopædia article
COAL PLANTS .
Carboniferous land plants, some of which were preserved in coal
balls , were very similar to those of the preceding Late
but new groups also appeared at this time. Ancient in situ
lycopsid , probably
Sigillaria , with attached stigmarian roots .
Base of a lycopsid showing connection with bifurcating stigmarian
The main Early
Carboniferous plants were the Equisetales
Sphenophyllales (scrambling plants),
Lepidodendrales (scale trees),
Medullosales (informally included in the "seed ferns ", an artificial
assemblage of a number of early gymnosperm groups) and the Cordaitales
. These continued to dominate throughout the period, but during late
Carboniferous , several other groups,
Cycadophyta (cycads), the
Callistophytales (another group of "seed ferns"), and the Voltziales
(related to and sometimes included under the conifers ), appeared.
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.
Lepidodendron (with its cone called
Anabathra , Lepidophloios and
Sigillaria . The roots of several of
these forms are known as
Stigmaria . Unlike present-day trees, their
secondary growth 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
The fronds of some
Carboniferous ferns are almost identical with
those of living species. Probably many species were epiphytic . Fossil
ferns and "seed ferns" include
Cyclopteris , Neuropteris
Alethopteris , and
Sphenopteris ; Megaphyton and Caulopteris were
Equisetales included the common giant form
Calamites , with a
trunk diameter of 30 to 60 cm (24 in) and a height of up to 20 m (66
Sphenophyllum was a slender climbing plant with whorls of leaves,
which was probably related both to the calamites and the lycopods.
Cordaites , a tall plant (6 to over 30 meters) with strap-like
leaves, was related to the cycads and conifers; the catkin -like
reproductive organs, which bore ovules/seeds, is called
These plants were thought to live in swamps. True coniferous trees
Walchia , of the order Voltziales) appear later in the Carboniferous,
and preferred higher drier ground.
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In the oceans the most important marine invertebrate groups are the
Foraminifera , corals ,
Ostracoda , brachiopods , ammonoids
, hederelloids , microconchids and echinoderms (especially crinoids ).
For the first time foraminifera take a prominent part in the marine
faunas. The large spindle-shaped genus Fusulina 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 .
The microscopic shells of radiolarians are found in cherts of this
age in the Culm of
Cornwall , and in Russia, Germany and
elsewhere. Sponges are known from spicules and anchor ropes, and
include various forms such as the Calcispongea Cotyliscus and
Girtycoelia, the demosponge Chaetetes, and the genus of unusual
colonial glass sponges
Both reef -building and solitary corals diversify and flourish; these
include both rugose (for example, Caninia , Corwenia, Neozaphrentis),
heterocorals, and tabulate (for example, Chladochonus, Michelinia)
Conularids were well represented by Conularia
Bryozoa are abundant in some regions; the fenestellids including
Fenestella, Polypora, and Archimedes , so named because it is in the
shape of an
Archimedean screw . Brachiopods are also abundant; they
include productids , some of which (for example,
reached very large (for brachiopods) size and had very thick shells,
while others like
Chonetes were more conservative in form. Athyridids
, spiriferids , rhynchonellids , and terebratulids are also very
common. Inarticulate forms include Discina and Crania . Some species
and genera had a very wide distribution with only minor variations.
Annelids such as Serpulites are common fossils in some horizons.
Among the mollusca, the bivalves continue to increase in numbers and
importance. Typical genera include
Carbonicola , Edmondia, and Modiola Gastropods are also
numerous, including the genera Murchisonia,
Euomphalus , Naticopsis.
Nautiloid cephalopods are represented by tightly coiled nautilids ,
with straight-shelled and curved-shelled forms becoming increasingly
Goniatite ammonoids are common.
Trilobites are rarer than in previous periods, on a steady trend
towards extinction, represented only by the proetid group.
a class of crustaceans , were abundant as representatives of the
meiobenthos ; genera included Amphissites, Bairdia, Beyrichiopsis,
Cavellina, Coryellina, Cribroconcha, Hollinella, Kirkbya, Knoxiella,
Amongst the echinoderms , 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 and Palaeechinus were
also present. The blastoids , 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.
Aviculopecten subcardiformis; a bivalve from the Logan Formation
(Lower Carboniferous) of
Wooster, Ohio (external mold).
Bivalves (Aviculopecten) and brachiopods (Syringothyris) in the Logan
Formation (Lower Carboniferous) in Wooster, Ohio.
Syringothyris sp.; a spiriferid brachiopod from the Logan Formation
(Lower Carboniferous) of
Wooster, Ohio (internal mold).
Palaeophycus ichnosp.; a trace fossil from the
Logan Formation (Lower
Carboniferous) of Wooster, Ohio.
Crinoid calyx from the Lower
Carboniferous of Ohio with a conical
platyceratid gastropod (Palaeocapulus acutirostre) attached.
Conulariid from the Lower
Carboniferous of Indiana.
Tabulate coral (a syringoporid); Boone
Carboniferous) near Hiwasse, Arkansas.
FRESHWATER AND LAGOONAL INVERTEBRATES
Carboniferous invertebrates include various bivalve
molluscs that lived in brackish or fresh water, such as Anthraconaia ,
Naiadites , and
Carbonicola ; diverse crustaceans such as
Carbonita , Darwinula , Estheria , Acanthocaris , Dithyrocaris , and
Anthrapalaemon . The upper
Carboniferous giant spider-like
Megarachne grew to legspans of 50 cm (20 in).
The Eurypterids were also diverse, and are represented by such genera
Megarachne (originally misinterpreted as a giant
spider, hence its name) and the specialised very large Hibbertopterus
. 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 being found in the thin beds known as marine bands.
Fossil remains of air-breathing insects , myriapods and arachnids
are known from the late Carboniferous, but so far not from the early
Carboniferous. The first true priapulids appeared during this period.
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 to grow
larger with the up to 2.6 metres long millipede-like Arthropleura
being the largest known land invertebrate of all time. Among the
insect groups are the huge predatory
Protodonata (griffinflies), among
Meganeura , a giant dragonfly -like insect and with a
wingspan of ca. 75 cm (30 in) — the largest flying insect ever to
roam the planet. Further groups are the Syntonopterodea (relatives of
present-day mayflies ), the abundant and often large sap-sucking
Palaeodictyopteroidea , the diverse herbivorous
Protorthoptera , and
Dictyoptera (ancestors of cockroaches ). Many insects
have been obtained from the coalfields of
and from the hollow trunks of fossil trees in Nova Scotia. Some
British coalfields have yielded good specimens: Archaeoptitus , from
the Derbyshire coalfield, had a spread of wing extending to more than
35 cm; some specimens (Brodia ) still exhibit traces of brilliant wing
colors. In the Nova Scotian tree trunks land snails (Archaeozonites ,
Dendropupa ) have been found.
Carboniferous giant dragonfly-like insect
Meganeura grew to
wingspans of 75 cm (30 in).
Pulmonoscorpius from the early
Carboniferous reached a
length of up to 70 cm (28 in).
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 ; some, the
petalodonts , 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 , the
Palaeonisciformes found in coastal
waters also appear to have migrated to rivers. Sarcopterygian fish
were also prominent, and one group, the Rhizodonts , reached very
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
Acanthodes , Cheirodus , and
Sharks (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
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 which possessed a flat brush-like dorsal fin with a
patch of denticles on its top.
Stethacanthus ' unusual fin may have
been used in mating rituals.
Akmonistion of the shark order
Symmoriida roamed the oceans of the
Falcatus was a
Carboniferous shark, with a high degree of sexual
Carboniferous amphibians 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
Labyrinthodontia . 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 15 cm (6
in) long. Some
Carboniferous amphibians were aquatic and lived in
Proterogyrinus ); others may have been
Hyloplesion ) or terrestrial
Carboniferous Rainforest Collapse slowed the evolution of
amphibians who could not survive as well in the cooler, drier
conditions. Reptiles, however, prospered due to specific key
adaptations. One of the greatest evolutionary innovations of the
Carboniferous was the amniote egg, which allowed the laying of eggs in
a dry environment, allowing for the further exploitation of the land
by certain tetrapods . These included the earliest sauropsid reptiles
Hylonomus ), and the earliest known synapsid (
Archaeothyris ). These
small lizard-like animals quickly gave rise to many descendants,
reptiles , birds , and mammals .
Reptiles underwent a major evolutionary radiation in response to the
drier climate that preceded the rainforest collapse. By the end of
Carboniferous period, amniotes had already diversified into a
number of groups, including protorothyridids , captorhinids ,
araeoscelids , and several families of pelycosaurs .
The amphibian -like
Pederpes , the most primitive Mississippian
Hylonomus , the earliest sauropsid reptile , appeared in the
Petrolacosaurus , the first diapsid reptile known, lived during the
Archaeothyris was a very early synapsid and the oldest known.
Because plants and animals were growing in size and abundance in this
time (for example,
Lepidodendron ), land fungi diversified further.
Marine fungi still occupied the oceans. All modern classes of fungi
were present in the Late
Carboniferous (Pennsylvanian Epoch).
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Main article: 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 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. The gap saw the demise of the
ichthyostegalian labyrinthodonts, and the rise of the more advanced
temnospondyl and reptiliomorphan amphibians that so typify the
Carboniferous terrestrial vertebrate fauna.
CARBONIFEROUS RAINFOREST COLLAPSE
Carboniferous Rainforest Collapse
Before the end of the
Carboniferous Period, an extinction event
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 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 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.
Carboniferous Rainforest Collapse
East Kirkton Quarry ; c. 350 mya;
Bathgate , Scotland
Hamilton Quarry ; 320 mya;
Kansas , US
Mazon Creek ; 300 mya;
Illinois , US
List of fossil sites
List of fossil sites (with link directory)
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