reached substantial diversity during this time, leading the Devonian to often be dubbed the Age of Fishes
. The placoderms
began dominating almost every known aquatic environment. The ancestors of all four-limbed vertebrates (tetrapods
) began adapting to walking on land, as their strong pectoral and pelvic fins gradually evolved into legs.
In the oceans, primitive sharks
became more numerous than in the Silurian and Late Ordovician
The rocks of Lummaton Quarry in Torquay
in Devon played an early role in defining the Devonian period.
The period is named after Devon
, a county in southwestern England, where a controversial argument in the 1830s over the age and structure of the rocks found distributed throughout the county was eventually resolved by the definition of the Devonian period in the geological timescale. The Great Devonian Controversy
was a long period of vigorous argument and counter-argument between the main protagonists of Roderick Murchison
with Adam Sedgwick
against Henry De la Beche
supported by George Bellas Greenough
. Murchison and Sedgwick won the debate and named the period they proposed as the Devonian System.[a]
In nineteenth-century texts the Devonian has been called the "Old Red Age", after the red and brown terrestrial deposits known in the United Kingdom as the Old Red Sandstone
in which early fossil discoveries were found. Another common term is "Age of the Fishes",
referring to the evolution of several major groups of fish
that took place during the period. Older literature on the Anglo-Welsh basin divides it into the Downtonian, Dittonian, Breconian, and Farlovian stages, the latter three of which are placed in the Devonian.
The Devonian has also erroneously been characterised as a "greenhouse age", due to sampling bias
: most of the early Devonian-age discoveries came from the strata
of western Europe
and eastern North America
, which at the time straddled the Equator
as part of the supercontinent of Euramerica where fossil
signatures of widespread reefs indicate tropical climates
that were warm and moderately humid but in fact the climate in the Devonian differed greatly during its epochs
and between geographic regions. For example, during the Early Devonian, arid conditions were prevalent through much of the world including Siberia, Australia, North America, and China, but Africa and South America
had a warm temperate climate
. In the Late Devonian, by contrast, arid conditions were less prevalent across the world and temperate climates
were more common.
The Devonian Period is formally broken into Early, Middle and Late subdivisions. The rocks corresponding to those epochs
are referred to as belonging to the Lower, Middle and Upper parts of the Devonian System.
The Early Devonian lasted from 419.2
± 2.8 to 393.3
± 2.5 and began with the Lochkovian
± 2.8 to 410.8
± 2.5, which was followed by the Pragian
± 2.8 to 407.6
± 2.5 and then by the Emsian
, which lasted until the Middle Devonian began, 393.3
± 2.7 million years ago.
During this time, the first ammonoids
appeared, descending from bactritoid nautiloids
. Ammonoids during this time period were simple and differed little from their nautiloid counterparts. These ammonoids belong to the order Agoniatitida
, which in later epochs evolved to new ammonoid orders, for example Goniatitida
. This class of cephalopod
molluscs would dominate the marine fauna until the beginning of the Mesozoic
The Middle Devonian comprised two subdivisions: first the Eifelian
, which then gave way to the Givetian 387.7
± 2.7 million years ago. During this time the jawless agnathan
fishes began to decline in diversity in freshwater and marine environments partly due to drastic environmental changes and partly due to the increasing competition, predation, and diversity of jawed fishes
. The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop such essential characteristics as well developed lungs, and the ability to crawl out of the water and onto the land for short periods of time.
Finally, the Late Devonian started with the Frasnian
± 2.8 to 372.2
± 2.5, during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing Famennian
subdivision, the beginning and end of which are marked with extinction events. This lasted until the end of the Devonian, 358.9
± 2.5 million years ago.
The Devonian was a relatively warm period, and probably lacked any glaciers
. The temperature gradient from the equator to the poles was not as large as it is today. The weather was also very arid, mostly along the equator where it was the driest.
Reconstruction of tropical sea surface temperature from conodont apatite
implies an average value of 30 °C (86 °F) in the Early Devonian.
levels dropped steeply throughout the Devonian period. The newly evolved forests drew carbon out of the atmosphere, which were then buried into sediments. This may be reflected by a Mid-Devonian cooling of around 5 °C (9 °F).
The Late Devonian warmed to levels equivalent to the Early Devonian; while there is no corresponding increase in CO2
concentrations, continental weathering increases (as predicted by warmer temperatures); further, a range of evidence, such as plant distribution, points to a Late Devonian warming.
The climate would have affected the dominant organisms in reefs
would have been the main reef-forming organisms in warm periods, with corals and stromatoporoid
sponges taking the dominant role in cooler times. The warming at the end of the Devonian may even have contributed to the extinction of the stromatoporoids.
The west coast of Devonian North America, by contrast, was a passive margin with deep silty embayments, river deltas and estuaries, found today in Idaho
; an approaching volcanic island arc
reached the steep slope of the continental shelf in Late Devonian times and began to uplift deep water deposits, a collision that was the prelude to the mountain-building episode at the beginning of the Carboniferous called the Antler orogeny
Spindle diagram for the evolution of vertebrates.
Sea levels in the Devonian were generally high. Marine faunas continued to be dominated by bryozoa
, diverse and abundant brachiopods
, the enigmatic hederellids
. Lily-like crinoids
(animals, their resemblance to flowers notwithstanding) were abundant, and trilobites
were still fairly common. Among vertebrates, jawless armored fish (ostracoderms
) declined in diversity, while the jawed fish (gnathostomes) simultaneously increased in both the sea and fresh water
. Armored placoderms were numerous during the lower stages of the Devonian Period and became extinct in the Late Devonian, perhaps because of competition for food against the other fish species. Early cartilaginous (Chondrichthyes
) and bony fishes (Osteichthyes
) also become diverse and played a large role within the Devonian seas. The first abundant genus of shark, Cladoselache
, appeared in the oceans during the Devonian Period. The great diversity of fish around at the time has led to the Devonian being given the name "The Age of Fish" in popular culture.
The first ammonites also appeared during or slightly before the early Devonian Period around 400 Mya.
A now dry barrier reef, located in present-day Kimberley Basin
of northwest Australia
, once extended 350 km (220 mi), fringing a Devonian continent.
Reefs in general are built by various carbonate
-secreting organisms that have the ability to erect wave-resistant structures close to sea level. Although modern reefs are constructed mainly by corals and calcareous algae
, Devonian reefs were either microbial reefs built up mostly by autotrophic cyanobacteria
, or coral-stromatoporoid reefs built up by coral-like stromatoporoids
and tabulate and rugose corals
. Microbial reefs dominated under the warmer conditions of the early and late Devonian, while coral-stromatoporoid reefs dominated during the cooler middle Devonian.
Prototaxites milwaukeensis, a large fungus, initially thought to be a marine alga, from the Middle Devonian of Wisconsin
By the Devonian Period, life was well underway in its colonisation of the land. The moss
forests and bacterial
and algal mats of the Silurian were joined early in the period by primitive rooted plants that created the first stable soils
and harbored arthropods like mites
(although arthropods appeared on land much earlier than in the Early Devonian
and the existence of fossils such as Protichnites
suggest that amphibious arthropods may have appeared as early as the Cambrian
). By far the largest land organism at the beginning of this period was the enigmatic Prototaxites
, which was possibly the fruiting body of an enormous fungus,
rolled liverwort mat,
or another organism of uncertain affinities
that stood more than 8 metres (26 ft) tall, and towered over the low, carpet-like vegetation during the early part of the Devonian. Also the first possible fossils of insects
appeared around 416 Mya, in the Early Devonian. Evidence for the earliest tetrapods
takes the form of trace fossils in shallow lagoon environments within a marine carbonate platform / shelf during the Middle Devonian,
although these traces have been questioned and an interpretation as fish feeding traces (Piscichnus
) has been advanced.
The greening of land
The Devonian period marks the beginning of extensive land colonisation by plants
. With large land-dwelling herbivores
not yet present, large forests grew and shaped the landscape.
Many Early Devonian plants
did not have true roots or leaves like extant plants although vascular tissue is observed in many of those plants. Some of the early land plants such as Drepanophycus
likely spread by vegetative growth and spores.
The earliest land plants such as Cooksonia
consisted of leafless, dichotomous
axes and terminal sporangia and were generally very short-statured, and grew hardly more than a few centimetres tall.
Fossils of Armoricaphyton chateaupannense
, about 400 million years old, represent the oldest known plants with woody
By the Middle Devonian, shrub-like forests of primitive plants existed: lycophytes
, and progymnosperms
had evolved. Most of these plants had true roots and leaves, and many were quite tall. The earliest-known trees appeared in the Middle Devonian
These included a lineage of lycopods and another arborescent, woody vascular plant, the cladoxylopsids
were able to grow to large size on dry land because they had evolved the ability to biosynthesize lignin
, which gave them physical rigidity and improved the effectiveness of their vascular system while giving them resistance to pathogens and herbivores.
These are the oldest-known trees of the world's first forests. By the end of the Devonian, the first seed-forming plants had appeared. This rapid appearance of so many plant groups and growth forms has been called the "Devonian Explosion".
Animals and the first soils
Primitive arthropods co-evolved with this diversified terrestrial vegetation structure. The evolving co-dependence of insects and seed-plants that characterised a recognisably modern world had its genesis in the Late Devonian period. The development of soils and plant root systems probably led to changes in the speed and pattern of erosion
and sediment deposition. The rapid evolution of a terrestrial ecosystem that contained copious animals opened the way for the first vertebrates
to seek out a terrestrial living. By the end of the Devonian, arthropods were solidly established on the land.
, one of the largest armoured fish ever to roam the planet, lived during the Late Devonian
Lower jaw of Eastmanosteus pustulosus from the Middle Devonian of Wisconsin
Tooth of the lobe-finned fish Onychodus from the Middle Devonian of Wisconsin
Melocrinites nodosus spinosus
, a spiny, stalked crinoid
from the Middle Devonian of Wisconsin
Enrolled phacopid trilobite
from the Devonian of Ohio
The common tabulate coral Aulopora
from the Middle Devonian of Ohio – view of colony encrusting a brachiopod
Tropidoleptus carinatus, an orthid brachiopod from the Middle Devonian of New York
SEM image of a hederelloid from the Devonian of Michigan (largest tube diameter is 0.75 mm)
Devonian spiriferid brachiopod from Ohio
which served as a host substrate for a colony of hederelloids
Late Devonian extinction
The Late Devonian is characterised by three episodes of extinction ("Late D")
The Late Devonian extinction is not a single event, but rather is a series of pulsed extinctions at the Givetian-Frasnian boundary, the Frasnian-Famennian boundary, and the Devonian-Carboniferous boundary.
Together, these are considered one of the "Big Five" mass extinctions in Earth's history.
The Devonian extinction crisis primarily affected the marine community, and selectively affected shallow warm-water organisms rather than cool-water organisms. The most important group to be affected by this extinction event were the reef-builders of the great Devonian reef systems.
Amongst the severely affected marine groups were the brachiopods, trilobites, ammonites, and acritarchs
, and the world saw the disappearance of an estimated 96% of vertebrates like conodonts
and bony fishes
, and all of the ostracoderms and placoderms.
Land plants as well as freshwater species, such as our tetrapod ancestors, were relatively unaffected by the Late Devonian extinction event (there is a counterargument that the Devonian extinctions nearly wiped out the tetrapods
The reasons for the Late Devonian extinctions are still unknown, and all explanations remain speculative.
Canadian paleontologist Digby McLaren
suggested in 1969 that the Devonian extinction events were caused by an asteroid impact. However, while there were Late Devonian collision events (see the Alamo bolide impact
), little evidence supports the existence of a large enough Devonian crater.
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". William Lonsdale stated that in December 1837 he had suggested the existence of a stratum between the Silurian and Carboniferous ones:
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